{"type": "root", "children": [{"type": "p", "children": [{"type": "t", "text": "\nGRHL2 is emerging as a master regulator of epithelial cell identity that plays a multifaceted role in modulating the epithelial‐to‐mesenchymal transition (EMT) and its reverse process, mesenchymal‐to‐epithelial transition (MET). Several studies have established that GRHL2 functions as a phenotypic stability factor by maintaining a hybrid epithelial/mesenchymal phenotype in cancer cells, thereby influencing collective cell migration, metastasis and tumor initiation. In breast, lung, ovarian and other cancers, GRHL2 suppresses key EMT drivers (such as ZEB1) and promotes the expression of epithelial markers including E-cadherin and CD24, which together are essential for preserving epithelial cell fate and tissue integrity."}, {"type": "fg", "children": [{"type": "fg_fs", "start_ref": "1", "end_ref": "6"}]}, {"type": "t", "text": "\n"}]}, {"type": "t", "text": "\n\n"}, {"type": "p", "children": [{"type": "t", "text": "\nIn addition to its role in directing cell plasticity, GRHL2 directly regulates genes needed for epithelial differentiation and intercellular adhesion. Experimental evidence demonstrates that GRHL2 binds to cis‐regulatory elements of crucial junctional components such as E-cadherin and claudin-4, thereby facilitating the formation of tight and adherens junctions in diverse tissues including the gut, lung and skin. GRHL2 also participates in the maintenance of telomerase activity by binding to the human telomerase reverse transcriptase (hTERT) promoter, which contributes to the regulation of cellular replicative lifespan."}, {"type": "fg", "children": [{"type": "fg_fs", "start_ref": "7", "end_ref": "11"}]}, {"type": "t", "text": "\n"}]}, {"type": "t", "text": "\n\n"}, {"type": "p", "children": [{"type": "t", "text": "\nThe contributions of GRHL2 in cancer are complex and appear context-dependent. In some settings, GRHL2 exerts oncogenic activities by enhancing cell proliferation, sustaining survival signals via pathways such as PI3K/Akt, and partnering with steroid receptors in hormone-dependent tumors, whereas in other contexts its suppression of EMT confers tumor-suppressive attributes by limiting invasion and metastasis. For example, GRHL2 has been implicated in modulating the epithelial phenotype in breast, gastric, oral and colorectal cancers, with its altered expression correlating with proliferative indices and chemoresistance."}, {"type": "fg", "children": [{"type": "fg_fs", "start_ref": "12", "end_ref": "18"}]}, {"type": "t", "text": "\n"}]}, {"type": "t", "text": "\n\n"}, {"type": "p", "children": [{"type": "t", "text": "\nBeyond its activities in epithelial biology and cancer, GRHL2 genetic alterations or dysregulation have been linked to developmental and degenerative disorders. Mutations or regulatory variants in GRHL2 are associated with sensorineural hearing loss, vestibular dysfunction and ocular anomalies such as posterior polymorphous corneal dystrophy. These observations extend GRHL2’s role from tumor biology to diverse aspects of organ development and homeostasis, highlighting its importance in tissue-specific gene regulation."}, {"type": "fg", "children": [{"type": "fg_fs", "start_ref": "19", "end_ref": "22"}]}, {"type": "t", "text": "\n"}]}, {"type": "t", "text": "\n\n"}, {"type": "p", "children": [{"type": "t", "text": "\nGRHL2 also engages in epigenetic and transcriptional regulatory networks that influence its functions. It interacts with chromatin modifiers and transcription factors (for instance, p63 and histone methyltransferases KMT2C/D) to govern gene expression programs critical for controlling EMT and mediating responses to external stimuli, including those from the immune system. Moreover, GRHL2-dependent regulation of microRNAs (like the miR-200 family and miR-194) further modulates its impact on epithelial differentiation and metabolic pathways, as illustrated in studies investigating natural killer cell-mediated cytotoxicity and non-alcoholic fatty liver disease."}, {"type": "fg", "children": [{"type": "fg_fs", "start_ref": "23", "end_ref": "29"}]}, {"type": "t", "text": "\n"}]}, {"type": "rg", "children": [{"type": "r", "ref": 1, "children": [{"type": "t", "text": "Mohit Kumar Jolly, Satyendra C Tripathi, Dongya Jia, et al. 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