MMP3 and rheumatoid arthritis: Both enforced expression of WAKMAR2 and the LLDT-8 treatment abolished beneficial effects of miR-4478 overexpression on proliferation and invasion of RA FLS, expression of PCNA and Cyclin D1, as well as the levels of MMP-3, IL-1, and IL-6 (Figures 7B–F, Supplementary Figures 4B,E).