{"type": "root", "children": [{"type": "p", "children": [{"type": "t", "text": "\nSOX9 is a master transcription factor that plays a seminal role in development, especially in chondrogenesis and cartilage formation. In early skeletogenesis, SOX9 is required for the sequential differentiation of chondrocytes, where it promotes cell survival, suppresses apoptosis, and transcriptionally activates cartilage‐specific matrix genes such as COL2A1, often by cooperating with cofactors including CBP/p300 and Smad3 in response to TGF‑β signals."}, {"type": "fg", "children": [{"type": "fg_fs", "start_ref": "1", "end_ref": "6"}]}, {"type": "t", "text": " Its dosage and precise regulation are crucial for normal skeletal development, as alterations in long‐range enhancers and mutations affecting SOX9’s dimerization or transcriptional activity lead to congenital malformations such as campomelic dysplasia and Pierre Robin sequence."}, {"type": "fg", "children": [{"type": "fg_fs", "start_ref": "7", "end_ref": "9"}]}, {"type": "t", "text": " Furthermore, SOX9 is central to sex determination as it mediates the SRY-induced commitment of bipotential gonads toward testis development."}, {"type": "fg", "children": [{"type": "fg_f", "ref": "10"}]}, {"type": "t", "text": ""}]}, {"type": "t", "text": "\n\n"}, {"type": "p", "children": [{"type": "t", "text": "\nIn addition to its developmental functions, aberrant expression or misregulation of SOX9 is implicated in multiple cancers and stem cell phenotypes. In several epithelial malignancies, including breast, prostate, colorectal, gastric, esophageal, lung, and glioblastoma, SOX9 supports tumor growth, invasion, and metastasis by sustaining stem/progenitor cell states and reactivating developmental gene networks. For instance, cooperative activity with factors such as Slug drives the mammary stem cell state and therapy resistance in breast cancer"}, {"type": "fg", "children": [{"type": "fg_f", "ref": "11"}]}, {"type": "t", "text": "; in prostate cancer, SOX9 acts as a critical downstream effector of oncogenic drivers (e.g., ERG) and modulates androgen receptor signaling to promote tumor proliferation."}, {"type": "fg", "children": [{"type": "fg_fs", "start_ref": "13", "end_ref": "16"}]}, {"type": "t", "text": " In colorectal and gastric cancers, SOX9 is overexpressed where it drives oncogenic transcriptional programs, influences epithelial-to-mesenchymal transition and collaborates with other factors to maintain cancer stem-like properties."}, {"type": "fg", "children": [{"type": "fg_f", "ref": "17"}]}, {"type": "t", "text": " Moreover, in hepatocellular carcinoma and other neuroectodermal tumors with NF1 alterations, elevated SOX9 is linked to increased proliferation and self-renewal of cancer stem cells."}, {"type": "fg", "children": [{"type": "fg_f", "ref": "19"}]}, {"type": "t", "text": ""}]}, {"type": "t", "text": "\n\n"}, {"type": "p", "children": [{"type": "t", "text": "\nSOX9 function is further refined by its interactions and regulatory inputs in specific cellular contexts. In articular and hypertrophic chondrocytes, its expression is modulated by microRNAs (e.g., miR-145) and hypoxic signaling via HIF-2α, reflecting a multilayered regulation that adjusts extracellular matrix synthesis and cellular differentiation."}, {"type": "fg", "children": [{"type": "fg_f", "ref": "21"}]}, {"type": "t", "text": " In the central nervous system, SOX9 serves as an almost exclusive nuclear marker of astrocytes, highlighting its role in neural cell identity."}, {"type": "fg", "children": [{"type": "fg_f", "ref": "23"}]}, {"type": "t", "text": " Additionally, regulatory complexes involving co-activators such as Med25 and ubiquitin-like modifiers modulate SOX9’s transcriptional activity in a tissue-specific manner, as exemplified by its role in regulating genes critical for palate formation."}, {"type": "fg", "children": [{"type": "fg_f", "ref": "24"}]}, {"type": "t", "text": " Collectively, these diverse studies position SOX9 as a pivotal regulator in both normal development and disease, with its context-dependent functions offering promising avenues for diagnostic and therapeutic applications."}, {"type": "fg", "children": [{"type": "fg_fs", "start_ref": "25", "end_ref": "28"}]}, {"type": "t", "text": ""}]}, {"type": "rg", "children": [{"type": "r", "ref": 1, "children": [{"type": "t", "text": "Haruhiko Akiyama, Marie-Christine Chaboissier, James F Martin, et al. 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