The brain of Drosophila is structurally altered by sensory stimuli that the flies receive during their adult life. Size and fiber number of the mushroom bodies, central complex, and optic lobes are influenced by social, spatial, or olfactory cues. Recently, the optic lobes have been shown to depend on the light regime that flies experience. Structural plasticity in the brain is thought to be a correlate of functional adaptations and long-term memory. We therefore extend our investigation of volume changes to the calyces of the mushroom bodies and the central complex. We show that rearing flies in constant light for 4 days increases the volume of both structures by up to 15% compared to rearing them in total darkness. Much of this difference develops during the first day. The effect of light is not hormonally mediated, as monocularly deprived flies develop a smaller ipsilateral calyx. Mutant analysis suggests that light generates its effects through known visual pathways. In contrast to the optic lobes, in the calyx and central complex structural changes can be linked to cAMP signaling, as in the mutants dunce z and amnesiac 1 no volume differences are observed. Surprisingly, the mutant rutabaga ~ shows a prominent light-dependent volume increase in the calyx and central complex, dissociating structural from behavioral plasticity. In complete darkness wild-type flies grow larger calyces under crowded conditions in their normal culture vials than if kept in small groups on fresh food. This stimulating effect of crowding is not 1Corresponding author. observed in any of the cAMP mutants, including rutabaga 1.