{"type": "root", "children": [{"type": "p", "children": [{"type": "t", "text": "\nTWIST2 is a highly conserved basic helix–loop–helix transcription factor that plays a critical role in embryogenesis and mesenchymal tissue development. Mutations in TWIST2 have been linked to congenital disorders such as Setleis syndrome, where homozygous nonsense mutations result in truncated, unstable proteins and lead to abnormal facial and skin development. These findings underscore its fundamental importance in craniofacial morphogenesis and connective tissue maintenance."}, {"type": "fg", "children": [{"type": "fg_fs", "start_ref": "1", "end_ref": "5"}]}, {"type": "t", "text": "\n"}]}, {"type": "t", "text": "\n\n"}, {"type": "p", "children": [{"type": "t", "text": "\nIn a wide spectrum of human cancers—including breast, cervical, hepatocellular, colorectal, head and neck, ovarian, and rhabdomyosarcomas—TWIST2 is frequently found to be overexpressed. Its upregulation drives the epithelial–mesenchymal transition (EMT) by repressing epithelial markers such as E-cadherin and inducing mesenchymal traits that promote increased cell migration, invasion, metastasis, and even vasculogenic mimicry. TWIST2 also contributes to the enrichment of cancer stem–like cell properties and helps tumor cells overcome failsafe programs like oncogene-induced senescence. In several clinical settings, high TWIST2 levels correlate with adverse pathological features and poor patient outcomes."}, {"type": "fg", "children": [{"type": "fg_f", "ref": "1"}, {"type": "fg_f", "ref": "3"}, {"type": "fg_fs", "start_ref": "6", "end_ref": "18"}]}, {"type": "t", "text": "\n"}]}, {"type": "t", "text": "\n\n"}, {"type": "p", "children": [{"type": "t", "text": "\nBeyond its roles in driving EMT and metastasis, TWIST2 has context-dependent regulatory functions in both normal and malignant hematopoietic as well as immune cells. In chronic lymphocytic leukemia and acute lymphoblastic leukemia, epigenetic silencing or altered regulation of TWIST2 impacts cell proliferation—partly through direct activation of cell cycle inhibitors like p21—implying a tumor‐suppressive facet in certain hematologic contexts."}, {"type": "fg", "children": [{"type": "fg_fs", "start_ref": "19", "end_ref": "21"}]}, {"type": "t", "text": " In parallel, TWIST2 modulates immune responses; for example, by participating in complexes that influence cytokine production in macrophages and by regulating apoptotic receptor expression (such as CD7) in T-cell lymphomas, it contributes to altered immune cell survival and functionality."}, {"type": "fg", "children": [{"type": "fg_fs", "start_ref": "22", "end_ref": "26"}]}, {"type": "t", "text": " Moreover, TWIST2’s expression can be modulated by noncoding RNAs, which further establishes a positive feedback loop that reinforces EMT and metastatic behavior in cancers such as gastric carcinoma."}, {"type": "fg", "children": [{"type": "fg_f", "ref": "27"}]}, {"type": "t", "text": " Additional studies have implicated TWIST2 in governing metabolic pathways within skeletal muscle and influencing tumor behavior via crosstalk with other transcriptional regulators."}, {"type": "fg", "children": [{"type": "fg_f", "ref": "11"}, {"type": "fg_fs", "start_ref": "28", "end_ref": "30"}]}, {"type": "t", "text": "\n"}]}, {"type": "rg", "children": [{"type": "r", "ref": 1, "children": [{"type": "t", "text": "Stéphane Ansieau, Jeremy Bastid, Agnès Doreau, et al. 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