Vascular networks constantly reorganize to optimize function. Some veins grow, others shrink and disappear. How can a local adaptation mechanism account for the plethora of vein dynamics observed? Here, we quantify network-wide vein dynamics in the slime mold Physarum polycephalum. We identify that flow shear stress is driving adaptation yet with a time delay. Vein fate, however, depends, beyond shear stress, on the vein's connections to the network and relative position. Finally, as network architecture constantly changes vein fate is dynamic driving overall network reorganization including avalanches of vein disappearance. Addition of external stimuli on top drives vein adaptation dynamics – imprinting the stimuli location into the network architecture. The memory of the stimulus location within the network architecture is retained during network reorganization and impacts overall network function.