{"type": "root", "children": [{"type": "p", "children": [{"type": "t", "text": "\nHuman DNA polymerase κ (POLK) is a specialized Y‐family enzyme that functions primarily in translesion synthesis (TLS) by bypassing a variety of DNA lesions. Biochemical and structural studies have revealed that POLK possesses unique features—including an “N‐clasp” that encircles DNA, a constrained active‐site cleft, and distinctive PCNA‐interaction motifs—which enable it to accurately incorporate the correct nucleotide opposite certain bulky adducts (such as benzo[a]pyrene–dG, N²‐adducts, thymine glycol, and oxidized lesions like 8‑oxoG), and to efficiently extend from mismatched primer termini generated by another polymerase. In vitro kinetics and crystallographic analyses show that POLK’s substrate preferences and its insertion/extension balance are tightly dictated by its active site geometry and interactions with accessory factors (e.g., hREV1), as well as the DNA minor groove–directed lesion bypass mechanism that it shares with other TLS polymerases."}, {"type": "fg", "children": [{"type": "fg_fs", "start_ref": "1", "end_ref": "15"}]}, {"type": "t", "text": "\n"}]}, {"type": "t", "text": "\n\n"}, {"type": "p", "children": [{"type": "t", "text": "\nBeyond its core TLS activity, POLK plays an important role in maintaining replication fork integrity and facilitating DNA damage responses. Its regulated recruitment to stalled forks is promoted by Rad18‐mediated PCNA monoubiquitination, which in turn coordinates polymerase exchange and the assembly of checkpoint complexes such as the 9–1–1 clamp. POLK thereby contributes to the synthesis of short DNA intermediates required for proper activation and recovery from S‐phase checkpoints. In addition, studies using both biochemical reconstitution and cellular models have linked POLK function to efficient extension of HR D‐loops and replication through structured regions like common fragile sites. Multiple investigations using kinetic assays, mutational analyses, and computational modeling further underscore POLK’s role in controlling genomic stability by preventing deleterious fork collapse and unchecked error‐prone synthesis."}, {"type": "fg", "children": [{"type": "fg_fs", "start_ref": "16", "end_ref": "28"}]}, {"type": "t", "text": "\n"}]}, {"type": "t", "text": "\n\n"}, {"type": "p", "children": [{"type": "t", "text": "\nIn the clinical context, POLK has emerged as a factor with significant implications in human disease. Its non‐canonical functions include a crucial role in hepatitis B virus cccDNA formation, while deregulated expression and promoter alterations have been associated with various cancers—including gliomas, lung, head and neck, and breast carcinomas—as well as prostate malignancies. Furthermore, specific POLK polymorphisms and tumor‐associated mutations have been correlated with altered lesion bypass activity, replication stress responses, and chemoresistance. Recent cryo–electron microscopy studies have also provided insights into how modifications (for example, PCNA ubiquitination) modulate POLK’s recruitment and activity, highlighting its potential as a target for therapeutic intervention."}, {"type": "fg", "children": [{"type": "fg_fs", "start_ref": "29", "end_ref": "38"}]}, {"type": "t", "text": "\n"}]}, {"type": "rg", "children": [{"type": "r", "ref": 1, "children": [{"type": "t", "text": "Eiji Ohashi, Yoshiki Murakumo, Naoko Kanjo, et al. 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