As invadopodia are distinct from membrane ruffles in being protrusive matrix-degrading structures on the ventral surface of cancer cells enriched with F-actin and cortactin in addition to the above-mentioned markers32, 33, we examined Agrin’s role in invadopodia formation by a gelatin-invadopodia assay simulating degradation of ECM. Here, AGRN is linked to cancer.