{"type": "root", "children": [{"type": "p", "children": [{"type": "t", "text": "\nDNA polymerase iota (POLI) is a member of the Y‐family of translesion synthesis (TLS) polymerases with unique biochemical and structural features. Its active site is highly specialized to promote Hoogsteen base pairing, which underpins its unusual template specificity—displaying high efficiency and fidelity when copying purines (especially adenine) while often being error‐prone opposite pyrimidines. In addition to its inherent polymerase activity, POLI exhibits deoxyribose phosphate lyase activity and can participate in base excision repair pathways. It is further equipped with evolutionarily conserved ubiquitin–binding motifs (UBMs) that facilitate its interaction with ubiquitinated proliferating cell nuclear antigen (PCNA), a modification critical for recruiting TLS polymerases to stalled replication forks and for orchestrating lesion bypass, as seen in somatic hypermutation processes."}, {"type": "fg", "children": [{"type": "fg_fs", "start_ref": "1", "end_ref": "10"}]}, {"type": "t", "text": "\n"}]}, {"type": "t", "text": "\n\n"}, {"type": "p", "children": [{"type": "t", "text": "\nPOLI’s recruitment to replication foci and its activity in TLS is tightly regulated by multiple protein–protein interactions and posttranslational modifications. Its ubiquitin–binding domains enable POLI to interact not only with monoubiquitinated PCNA but also directly with ubiquitin chains—features that help control its spatial and temporal engagement at stalled forks. Moreover, specific sequence variants (SNPs) in the POLI gene have been linked to altered DNA repair efficiency and increased susceptibility to cancers such as lung adenocarcinoma and squamous cell carcinoma. POLI is highly dynamic within the nucleus, transiently probing DNA and collaborating with other TLS polymerases (for example, POLη) to ensure controlled bypass of a variety of DNA lesions."}, {"type": "fg", "children": [{"type": "fg_fs", "start_ref": "11", "end_ref": "20"}]}, {"type": "t", "text": "\n"}]}, {"type": "t", "text": "\n\n"}, {"type": "p", "children": [{"type": "t", "text": "\nBeyond its classic role in lesion bypass, POLI is implicated in broader genomic maintenance and cancer-relevant processes. Its activity contributes to highly controlled TLS during replication stress and is integrated into p53-dependent DNA damage tolerance pathways, wherein POLI complexes help decelerate nascent DNA elongation to promote safe, homology-directed bypass of stalled forks. Dysregulation of POLI—in part through altered expression, aberrant ubiquitination, or genetic variations—has been linked to hypermutagenesis in breast, esophageal, and bladder cancers, as well as to synthetic lethal interactions in cells deficient for Fanconi anemia genes. Furthermore, emerging findings indicate that POLI participates in modulating replication dynamics in stem cells, ensuring a balance between replication speed and genome integrity."}, {"type": "fg", "children": [{"type": "fg_fs", "start_ref": "21", "end_ref": "38"}]}, {"type": "t", "text": "\n"}]}, {"type": "rg", "children": [{"type": "r", "ref": 1, "children": [{"type": "t", "text": "Ahmad Faili, Said Aoufouchi, Eric Flatter, et al. 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