AbstractTo navigate towards odor in a turbulent environment, animals integrate odor value cues with wind direction cues. The central neural circuits subserving this behavior are unknown. Here we used optogenetic activation to identify neurons in the lateral horn (LH), mushroom body (MB), and fan-shaped body (FB), that drive upwind orientation in walkingDrosophila. Using calcium imaging, we show that MB/LH neurons encode odor independent of wind direction, and that odor and wind are integrated within hΔC local neurons of the FB to drive re-orientation. Based on connectome data, we model an FB circuit that allows odor to switch behavioral orientation to wind. Our work identifies central neural circuits that integrate value and direction signals to generate an essential goal-directed behavior.