Hierarchical control over foraging behavior by anterior cingulate cortex

Foraging is a natural behavior that involves making sequential decisions to maximize rewards while minimizing the costs incurred when doing so. The prevalence of foraging across species suggests that a common brain computation underlies its implementation. Although anterior cingulate cortex is believed to contribute to foraging behavior, its specific role has been contentious, with predominant theories arguing either that it encodes environmental value or choice difficulty. Additionally, recent attempts to characterize foraging have taken place within the reinforcement learning framework, with increasingly complex models scaling with task complexity. Here we review reinforcement learning foraging models, highlighting the hierarchical structure of many foraging problems. We extend this literature by proposing that ACC guides foraging according to principles of modelbased hierarchical reinforcement learning. This idea holds that ACC function is organized hierarchically along a rostral-caudal gradient, with rostral structures monitoring the status and completion of high-level task goals (like finding food), and midcingulate structures overseeing the execution of task options (subgoals, like harvesting fruit) and lower -level actions (such as grabbing an apple).