Additionally, paired single-cell and transposase-accessible chromatin sequencing (ATAC-seq) analysis of synovial biopsies from RA patients, combined with multiplex imaging and spatial transcriptomics, suggested that local TNF, IFN-γ or IL-1β exposure can drive FLS heterogeneity, and also indicated that functional gene expression programs in fibroblasts may be shared across tissues and diseases (45). Here, TNF is linked to rheumatoid arthritis.