“…In vivo, cells crawl through topographically intricate environments, such as the extracellular matrix, blood and lymphatic vessels, other cells, etc., that can significantly influence migration strategies [6][7][8][9]. For instance, it has been shown that local anisotropy in the underlying substrate, in the form of adhesive ratchets [10][11][12] or three-dimensional structures on the subcellular scale [10,13,14], can lead to directed motion even in the absence of chemical stimuli. More recently, Wondergem and coworkers demonstrated directed migration of single cells using a spatial gradient in the density of cell-sized topographical features [15].…”