{"type": "root", "children": [{"type": "p", "children": [{"type": "t", "text": "\nTestin (TES) is a multifunctional scaffold protein that localizes prominently at focal adhesions and cell–cell contacts, where it plays a central role in organizing the actin cytoskeleton and regulating cell motility. Structural studies have revealed that TES’s LIM domains—particularly LIM3—mediate binding to actin‐regulatory proteins such as Mena by occluding the FPPPP‐binding interface of EVH1 domains, thereby competing with other cytoskeletal ligands and modulating their localization at the cell leading edge and focal adhesions."}, {"type": "fg", "children": [{"type": "fg_f", "ref": "1"}]}, {"type": "t", "text": " Moreover, TES is recruited to specialized focal adherens junctions in a mechanosensitive manner alongside VASP and zyxin, indicating that it contributes to force‐dependent regulation of cell–cell contacts."}, {"type": "fg", "children": [{"type": "fg_f", "ref": "2"}]}, {"type": "t", "text": " In addition, RNAi‐mediated knockdown studies have demonstrated that loss of TES from focal adhesions disrupts actin stress fibre formation and reduces RhoA activity"}, {"type": "fg", "children": [{"type": "fg_f", "ref": "3"}]}, {"type": "t", "text": ", while crystallographic analyses further detail its interaction with actin‐related proteins such as Arp7A and reveal that TES can adopt distinct conformational states through intramolecular PET and LIM domain interactions."}, {"type": "fg", "children": [{"type": "fg_f", "ref": "4"}]}, {"type": "t", "text": " Complementary studies have also shown that TES regulates cytoskeletal dynamics by interacting with the calcium‐sensing receptor to potentiate RhoA signalling."}, {"type": "fg", "children": [{"type": "fg_f", "ref": "6"}]}, {"type": "t", "text": "\n"}]}, {"type": "t", "text": "\n\n"}, {"type": "p", "children": [{"type": "t", "text": "\nIn a broad spectrum of malignancies—including prostate, breast, gastric, head and neck, endometrial, and lung cancers—TES has emerged as a potent tumor suppressor. Reduced TES expression, frequently resulting from loss of heterozygosity, promoter hypermethylation, or accelerated proteasomal degradation, correlates with enhanced cell proliferation, migration, invasion, and epithelial–mesenchymal transition (EMT). For instance, in prostate cancers, diminished TES mRNA levels align with LOH at the 7q31 locus"}, {"type": "fg", "children": [{"type": "fg_f", "ref": "7"}]}, {"type": "t", "text": ", whereas in breast and gastric carcinomas, progressive loss of TES is associated with increased invasive and angiogenic potential."}, {"type": "fg", "children": [{"type": "fg_f", "ref": "8"}]}, {"type": "t", "text": " TES downregulation has also been documented in head and neck cancers"}, {"type": "fg", "children": [{"type": "fg_f", "ref": "10"}]}, {"type": "t", "text": "and is implicated in promoting a more aggressive phenotype via hypermethylation and proteolytic degradation in gastric cancers."}, {"type": "fg", "children": [{"type": "fg_f", "ref": "11"}]}, {"type": "t", "text": " Furthermore, studies in endometrial"}, {"type": "fg", "children": [{"type": "fg_f", "ref": "12"}]}, {"type": "t", "text": ", breast"}, {"type": "fg", "children": [{"type": "fg_f", "ref": "13"}]}, {"type": "t", "text": ", and childhood acute lymphoblastic leukemia"}, {"type": "fg", "children": [{"type": "fg_f", "ref": "14"}]}, {"type": "t", "text": "illustrate that TES silencing is linked to EMT and poor clinical outcomes. TES re-expression in non‐small cell lung cancer models"}, {"type": "fg", "children": [{"type": "fg_f", "ref": "15"}]}, {"type": "t", "text": "and gastric cancers"}, {"type": "fg", "children": [{"type": "fg_f", "ref": "16"}]}, {"type": "t", "text": "has been shown to restrain tumor growth and invasiveness, while further studies in breast cancer demonstrate that loss of TES facilitates reduced cellular adhesion and enhanced metastatic behavior through modulation of RhoA signalling."}, {"type": "fg", "children": [{"type": "fg_f", "ref": "17"}]}, {"type": "t", "text": " Clinical correlations also indicate that lower TES levels predict worse prognosis in gastric"}, {"type": "fg", "children": [{"type": "fg_f", "ref": "18"}]}, {"type": "t", "text": "and melanoma"}, {"type": "fg", "children": [{"type": "fg_f", "ref": "19"}]}, {"type": "t", "text": "patients, and genetic association studies in prostate cancer further support its tumor‐suppressive role."}, {"type": "fg", "children": [{"type": "fg_f", "ref": "20"}]}, {"type": "t", "text": " Recent clinical evaluations in lung cancer also underscore the prognostic significance of TES expression."}, {"type": "fg", "children": [{"type": "fg_f", "ref": "21"}]}, {"type": "t", "text": "\n"}]}, {"type": "t", "text": "\n\n"}, {"type": "p", "children": [{"type": "t", "text": "\nBeyond its established functions in tumor suppression and cytoskeletal regulation, TES is emerging as an important player in other pathophysiological contexts. Biochemical investigations have revealed that TES adopts multiple conformational states and can localize to unexpected compartments such as the nucleolus, suggesting a versatility in its functional roles."}, {"type": "fg", "children": [{"type": "fg_f", "ref": "22"}]}, {"type": "t", "text": " Proteomic analyses have expanded the known TES interactome to include novel partners like TGFB1I1 and a short form of the glucocorticoid receptor, linking TES to the modulation of cell spreading and additional signal transduction pathways."}, {"type": "fg", "children": [{"type": "fg_f", "ref": "23"}]}, {"type": "t", "text": " Its regulatory influence extends to nasopharyngeal carcinoma, where TES modulates proliferation and migration"}, {"type": "fg", "children": [{"type": "fg_f", "ref": "24"}]}, {"type": "t", "text": ", and studies in colorectal cancer have identified TES as a key component of cell–cell junctions, with its loss correlating with increased cell motility and proliferation."}, {"type": "fg", "children": [{"type": "fg_f", "ref": "25"}]}, {"type": "t", "text": " Intriguingly, in cardiovascular research, TES has been shown to counteract cardiac hypertrophy by interacting with calcineurin and attenuating downstream hypertrophic signalling."}, {"type": "fg", "children": [{"type": "fg_f", "ref": "26"}]}, {"type": "t", "text": " Collectively, these findings underscore TES’s pivotal roles as a mechanosensitive cytoskeletal regulator and a tumor suppressor, with broader implications for cellular homeostasis and disease.\n"}]}, {"type": "rg", "children": [{"type": "r", "ref": 1, "children": [{"type": "t", "text": "Batiste Boëda, David C Briggs, Theresa Higgins, et al. 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