(A) Schematic model showing how an animal’s context, including whether it is walking spontaneously in clean air or engaged in active odor pursuit, the navigational strategy it employs, and its satiety state, shapes the moment-to-moment relationships between DAN activity and different behavioral variables (grey dial), giving rise to the longer timescale relationships in which DAN activity is preferentially tuned to the motor actions that subserve odor pursuit. An animal’s context also coordinately influences behavior (thick grey arrows). Acute manipulation of DAN activity alters behavior, highlighting how the mushroom body dopaminergic system is embedded within a larger feedback loop (grey dashed arrow). (B) Schematic depicting how the model in (A) produces the behavior and neural activity we observe in the low (left) and high (right) airflow contexts. Under low airflow conditions (left), the γ4-|heading| relationship is selectively strengthened. When a fly encounters the odor, it reorients and elevated γ4 DAN activity promotes upwind tracking towards the odor source. Conversely, under high airflow conditions (right), the γ4 DAN activity-forward velocity relationship is selectively strengthened. When a fly encounters the odor, increased γ4 DAN activity also promotes upwind tracking towards the odor source despite a fly using different actions for pursuit.