The EMT transcription factor Twist is reported to induce the formation of actin-enriched membrane protrusions called aggressive pseudopods (invasive pseudopods are a special actin-based membrane protrusion found in cancer cells, and degrade ECM by targeting proteases); aggressive pseudopodia can recruit MMP-2, MMP-9, and MMP-14 to the leading edge where they degrade collagen and the basement membrane ECM, thereby promoting tumor invasion and metastasis [199, 200]. Here, MMP2 is linked to neoplasm.