WAKMAR2 silencing with the presence of LLDT-8 recovered the 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).These results demonstrated the essential role of LLDT-8/WAKMAR2/miR-4478/E2F1/p53 axis in RA FLS (Figure 8). Here, E2F1 is linked to rheumatoid arthritis.