{"type": "root", "children": [{"type": "p", "children": [{"type": "t", "text": "\nZIC2 is a zinc‐finger transcription factor that plays a critical role in embryonic neural development and forebrain patterning. It regulates retinal ganglion cell axon guidance—mediating midline repulsion essential for proper binocular vision—and is pivotal for the formation of the forebrain, as evidenced by its association with holoprosencephaly and neural tube defects. Mutations or dosage alterations in ZIC2 have been shown in both human patients and animal models to result in a spectrum of midline anomalies, ranging from severe brain malformations to subtle craniofacial dysmorphisms. Moreover, studies indicate that ZIC2 functions via dual transcriptional mechanisms, containing distinct activation and repression domains, and interacts with key developmental signaling pathways (e.g., NODAL and β–catenin–TCF4) necessary for left–right axis specification and proper neural tube closure."}, {"type": "fg", "children": [{"type": "fg_fs", "start_ref": "1", "end_ref": "13"}]}, {"type": "t", "text": "\n"}]}, {"type": "t", "text": "\n\n"}, {"type": "p", "children": [{"type": "t", "text": "\nIn addition to its developmental functions, aberrant expression and dysregulation of ZIC2 have emerged as important contributors to tumorigenesis. Upregulated ZIC2 expression is frequently observed in various malignancies—including hepatocellular carcinoma, oral squamous cell carcinoma, pancreatic ductal adenocarcinoma, and ovarian, breast, and nasopharyngeal cancers—where it promotes cancer stem cell self‐renewal, cell proliferation, migration, and invasion. In these contexts, ZIC2 acts upstream of key oncogenic factors (for example, OCT4, PAK4, and STAT3) and modulates critical pathways such as the Wnt/β–catenin, Hedgehog, and TGF–β signaling cascades. Such activities underscore its utility not only as a prognostic marker but also as a potential therapeutic target, with its interactions often being modulated by microRNAs that either suppress or enhance its oncogenic effects."}, {"type": "fg", "children": [{"type": "fg_fs", "start_ref": "14", "end_ref": "27"}]}, {"type": "t", "text": "\n"}]}, {"type": "t", "text": "\n\n"}, {"type": "p", "children": [{"type": "t", "text": "\nRecent findings further expand the diverse regulatory repertoire of ZIC2 by implicating it in the control of enhancer activation and epigenetic remodeling during lineage specification and maintenance of pluripotency. In certain cancers such as colon and lung adenocarcinoma, ZIC2’s context‐dependent functions—mediated through its interaction with non–coding RNAs and recruitment of chromatin remodeling complexes—contribute to the fine‐tuning of gene expression programs that govern both tumor aggression and stem cell characteristics. These advanced insights into ZIC2’s multilevel regulatory mechanisms provide new avenues for targeted interventions in developmental disorders and cancer."}, {"type": "fg", "children": [{"type": "fg_fs", "start_ref": "28", "end_ref": "31"}]}, {"type": "t", "text": "\n"}]}, {"type": "rg", "children": [{"type": "r", "ref": 1, "children": [{"type": "t", "text": "Pascale Marcorelles, Philippe Loget, Catherine Fallet-Bianco, et al. 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