{"type": "root", "children": [{"type": "p", "children": [{"type": "t", "text": "\nmiR‐491—predominantly the 5p form and also its 3p counterpart—has emerged as a pivotal tumor suppressor across a broad spectrum of solid tumors. In oral squamous cell carcinoma, reduced miR‐491 expression correlates with poor overall survival, while its re‐expression inhibits cell migration, invasion, and lung metastasis via direct targeting of GIT1."}, {"type": "fg", "children": [{"type": "fg_fs", "start_ref": "1", "end_ref": "3"}]}, {"type": "t", "text": " In colorectal cancer, ectopic miR‐491 expression induces apoptosis through downregulation of the anti‐apoptotic protein Bcl‑X(L)"}, {"type": "fg", "children": [{"type": "fg_f", "ref": "4"}]}, {"type": "t", "text": ", and in ovarian cancer it similarly promotes apoptosis and overcomes anti‐apoptotic signals by targeting Bcl‑X(L) and epidermal growth factor receptor (EGFR)."}, {"type": "fg", "children": [{"type": "fg_f", "ref": "5"}]}, {"type": "t", "text": " Furthermore, in gastric cancer miR‑491 directly targets key effectors such as SNAIL and Wnt3a to suppress epithelial–mesenchymal transition and metastasis"}, {"type": "fg", "children": [{"type": "fg_f", "ref": "6"}]}, {"type": "t", "text": ", and in bladder cancer its tumor‐suppressive impact is evidenced by circRNA studies that show sponging of miR‑491 leads to de‐repression of MMP9."}, {"type": "fg", "children": [{"type": "fg_f", "ref": "8"}]}, {"type": "t", "text": "\n"}]}, {"type": "t", "text": "\n\n"}, {"type": "p", "children": [{"type": "t", "text": "\nIn addition to its role in curbing tumor aggressiveness, miR‐491 critically modulates chemoresistance and cancer stem cell properties. In hepatoma cells, for instance, miR‑491 suppresses the PI3K/Akt signalling cascade, thereby influencing HCV replication dynamics."}, {"type": "fg", "children": [{"type": "fg_f", "ref": "9"}]}, {"type": "t", "text": " In glioblastoma and estrogen receptor–positive breast cancer, miR‑491 (via both its arms) directly targets oncogenic drivers such as EGFR, CDK6, and the histone demethylase JMJD2B to inhibit proliferation and invasion."}, {"type": "fg", "children": [{"type": "fg_f", "ref": "10"}]}, {"type": "t", "text": " Its reduced levels have also been associated with disease progression in esophageal cancer (where multiple miRNAs are dysregulated)"}, {"type": "fg", "children": [{"type": "fg_f", "ref": "12"}]}, {"type": "t", "text": ", nasopharyngeal carcinoma where downregulation leads to derepression of Notch3"}, {"type": "fg", "children": [{"type": "fg_f", "ref": "13"}]}, {"type": "t", "text": ", and in dopaminergic systems where miR‑491, together with miR‑137, regulates dopamine transporter expression in a VNTR‐dependent manner."}, {"type": "fg", "children": [{"type": "fg_f", "ref": "14"}]}, {"type": "t", "text": " In tongue cancer, reduced miR‑491 levels confer chemoresistance by relieving inhibition of mTORC2 components"}, {"type": "fg", "children": [{"type": "fg_f", "ref": "15"}]}, {"type": "t", "text": ", and in hepatocellular and other cancers, modulation of miR‑491 impacts multidrug resistance via downregulation of ABCB1 and its transcriptional regulator Sp3."}, {"type": "fg", "children": [{"type": "fg_f", "ref": "17"}]}, {"type": "t", "text": " Moreover, genetic studies highlight that polymorphisms affecting miR‑491 binding—for example in the 3′‑UTR of MMP9—can further alter its regulatory efficiency"}, {"type": "fg", "children": [{"type": "fg_f", "ref": "19"}]}, {"type": "t", "text": ", while aspirin sensitizes hepatocellular carcinoma side‐population cells to doxorubicin via a miR‑491/ABCG2 axis."}, {"type": "fg", "children": [{"type": "fg_f", "ref": "20"}]}, {"type": "t", "text": " In cervical cancer, oncogenic JMJD2A represses miR‑491 expression, thereby promoting cell proliferation and survival, and in esophageal as well as other malignancies, diminished miR‑491 contributes to aggressive tumor behavior and therapeutic resistance."}, {"type": "fg", "children": [{"type": "fg_fs", "start_ref": "21", "end_ref": "26"}]}, {"type": "t", "text": "\n"}]}, {"type": "t", "text": "\n\n"}, {"type": "p", "children": [{"type": "t", "text": "\nA further layer of complexity in miR‑491’s function arises from its regulation by competing endogenous RNAs. Long noncoding RNAs such as XIST, MALAT1, and LINC00460—and circular RNAs including circ_0000212 and circ_0006988—act as molecular sponges that sequester miR‑491, abrogating its ability to repress targets like FOXP4, MAP3K3, and UBE2C; these interactions have been implicated in the progression of nasopharyngeal carcinoma, non–small cell lung cancer, colorectal cancer, and pancreatic ductal adenocarcinoma."}, {"type": "fg", "children": [{"type": "fg_fs", "start_ref": "27", "end_ref": "34"}]}, {"type": "t", "text": " Collectively, these studies underscore that miR‑491 is a central regulator of key signaling cascades—including the PI3K/Akt, NF‑κB, and Wnt/β‑catenin pathways—that govern apoptosis, proliferation, migration, metastasis, and chemoresistance in cancer.\n"}]}, {"type": "rg", "children": [{"type": "r", "ref": 1, "children": [{"type": "t", "text": "Haruo Nakano, Tatsuya Miyazawa, Keita Kinoshita, et al. 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