Insights from the Evolving Model of Two Cortical Visual Pathways

The two cortical visual pathways framework has had a profound influence on theories and empirical studies of the visual system for over 40 years. By grounding physiological responses and behavior in neuroanatomy, the framework provided a critical guide for understanding vision. Although the framework has evolved over time, as our understanding of the physiology and neuroanatomy expanded, cortical visual processing is still often conceptualized as two separate pathways emerging from the primary visual cortex that support distinct behaviors ("what" vs. "where/how"). Here, we take a historical perspective and review the continuing evolution of the framework, discussing key and often overlooked insights. Rather than a functional and neuroanatomical bifurcation into two independent serial, hierarchical pathways, the current evidence points to two highly recurrent heterarchies with heterogeneous connections to cortical regions and subcortical structures that flexibly support a wide variety of behaviors. Although many of the simplifying assumptions of the framework are belied by the evidence gathered since its initial proposal, the core insight of grounding function and behavior in neuroanatomy remains fundamental. Given this perspective, we highlight critical open questions and the need for a better understanding of neuroanatomy, particularly in the human.