E2F1 and rheumatoid arthritis: WAKMAR2 silencing in the presence of LLDT-8 released the inhibitory effects of miR-4478 on E2F1 and p53expression and recovered the beneficial effects of miR-4478 on RA FLS, suggesting that WAKMAR2/miR-4478/E2F1/p53 axis is essential for the action of LLDT-8 on RA FLS.