{"type": "root", "children": [{"type": "p", "children": [{"type": "t", "text": "\nP4HA1 is the essential catalytic subunit of collagen prolyl 4‐hydroxylase, the enzyme responsible for converting specific proline residues in protocollagen chains into (2S,4R)‐4-hydroxyproline. This posttranslational modification is critical for the proper folding, triple-helical stability, and secretion of collagen and thus for maintaining the structural integrity of the extracellular matrix (ECM). Multiple studies have demonstrated that P4HA1 expression is tightly regulated—via transcriptional mechanisms (including USF1/USF2 interactions) and by posttranscriptional modulators such as RNA-binding proteins and changes in mRNA stability—and that its activity is sensitive to environmental cues including oxygen tension and iron levels. These regulatory features underpin its role in tissue remodeling under both physiologic and pathologic conditions."}, {"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": "\nBeyond its canonical role in collagen biogenesis, aberrant P4HA1 expression has emerged as a critical driver of tumor progression and aggressiveness. Elevated levels of P4HA1 have been associated with increased cancer cell stemness, enhanced invasion, epithelial–mesenchymal transition (EMT), metabolic reprogramming, and chemoresistance across a variety of tumor types—including breast, prostate, ovarian, pancreatic, lung, colorectal, melanoma, and esophageal cancers. In these contexts, P4HA1 contributes not only by promoting ECM remodeling and collagen deposition but also by modulating the stability of hypoxia-inducible factor 1α (HIF-1α), thereby reinforcing a hypoxic, pro-tumorigenic microenvironment. Moreover, its expression is subject to negative regulation by tumor-suppressive microRNAs (such as miR-122 and miR-30e), highlighting its potential as a therapeutic target and prognostic biomarker."}, {"type": "fg", "children": [{"type": "fg_fs", "start_ref": "11", "end_ref": "25"}]}, {"type": "t", "text": "\n"}]}, {"type": "t", "text": "\n\n"}, {"type": "p", "children": [{"type": "t", "text": "\nAdditional studies have extended the impact of P4HA1 beyond cancer cell–intrinsic activities to include roles in vascular remodeling, fibrotic processes, and the regulation of cell fate. Altered P4HA1 expression has been implicated in compromising the stability of vascular basement membranes, promoting arterial wall remodeling, and even influencing resistance to ferroptosis via modulation of metabolic pathways such as the mevalonate pathway. Furthermore, in malignant gliomas and hypopharyngeal tumors, P4HA1 contributes to neovascularization and is emerging as a promising biomarker in non‐oncologic as well as oncologic settings. These findings collectively suggest that targeting P4HA1 may offer therapeutic benefits across multiple pathological conditions."}, {"type": "fg", "children": [{"type": "fg_fs", "start_ref": "26", "end_ref": "30"}]}, {"type": "t", "text": "\n"}]}, {"type": "rg", "children": [{"type": "r", "ref": 1, "children": [{"type": "t", "text": "Michael Fähling, Ralf Mrowka, Andreas Steege, et al. 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