Constructed with a substrate sensitive to MT1-MMP, the first MT1-MMP FRET sensor observed epidermal growth factor (EGF)-induced, cytoskeleton-dependent MT1-MMP activity at the leading edge of migrating cancer cells, which can be abolished by tissue inhibitor of metalloproteinase-2 (TIMP-2) (Ouyang et al., 2008a). This evidence concerns the gene MMP14 and cancer.