{"type": "root", "children": [{"type": "p", "children": [{"type": "t", "text": "\nPRODH encodes proline oxidase (POX), a mitochondrial enzyme that catalyzes the rate‐limiting first step in proline degradation. In numerous cancer models, PRODH/POX plays a dual role by not only channeling proline catabolism to generate ATP under nutrient stress but also by producing reactive oxygen species (ROS) that trigger apoptosis. For example, in pancreatic ductal adenocarcinoma cells, ingestion of collagen‐derived proline sustains metabolism when other fuels are scarce, and POX activity is essential for cell proliferation."}, {"type": "fg", "children": [{"type": "fg_f", "ref": "1"}]}, {"type": "t", "text": " Similarly, inducible expression of POX in colorectal and lung carcinoma cells activates both intrinsic mitochondrial and extrinsic death receptor pathways via ROS generation, modulating calcineurin/NFAT and MEK/ERK cascades while suppressing proliferative signaling (e.g., COX‐2, EGFR and Wnt/β‐catenin)."}, {"type": "fg", "children": [{"type": "fg_fs", "start_ref": "2", "end_ref": "7"}]}, {"type": "t", "text": " In some tumors, such as renal carcinoma, upregulation of miR‑23b directly targets and downregulates PRODH, thereby diminishing its tumor‐suppressive functions."}, {"type": "fg", "children": [{"type": "fg_f", "ref": "8"}]}, {"type": "t", "text": " Moreover, studies in cancer cell models have shown that under metabolic or oxidative stress, PRODH‐dependent ROS production can also drive autophagy, and that impaired proline availability compromises POX‐mediated apoptosis."}, {"type": "fg", "children": [{"type": "fg_f", "ref": "9"}]}, {"type": "t", "text": "\n"}]}, {"type": "t", "text": "\n\n"}, {"type": "p", "children": [{"type": "t", "text": "\nIn the nervous system, genetic alterations at the PRODH locus have been implicated in several neuropsychiatric conditions. Heterozygous deletions and missense mutations in PRODH lead to hyperprolinemia and are associated with an increased risk for schizophrenia, schizoaffective disorder, and autism spectrum disorders—findings that are especially notable in patients with 22q11.2 deletion syndrome. Disrupted PRODH activity is thought to affect neurotransmitter balance and synaptic signaling, contributing to altered brain structure and connectivity. Indeed, several studies have linked specific risk and protective PRODH alleles, alone or in concert with variation in COMT, to deficits in executive function, cognitive performance, and abnormal neural circuit organization."}, {"type": "fg", "children": [{"type": "fg_fs", "start_ref": "11", "end_ref": "17"}]}, {"type": "t", "text": " In addition, a human‐specific endogenous retroviral enhancer (hsERVPRODH) has been shown to synergize with cis‑regulatory elements to drive PRODH expression in the central nervous system, further underscoring its relevance in neurodevelopment."}, {"type": "fg", "children": [{"type": "fg_f", "ref": "18"}]}, {"type": "t", "text": " Imaging and genetic studies in subjects with 22q11.2 deletion syndrome have also reported that PRODH variants contribute to altered brain morphology and connectivity, while other investigations have associated low‐activity PRODH alleles with autism‐like features and cognitive impairment."}, {"type": "fg", "children": [{"type": "fg_fs", "start_ref": "19", "end_ref": "21"}]}, {"type": "t", "text": " Mutational analyses in pediatric hyperprolinemia further reveal that severe impairment of PRODH enzymatic function correlates with early‐onset neurodevelopmental deficits"}, {"type": "fg", "children": [{"type": "fg_f", "ref": "22"}]}, {"type": "t", "text": ", and comprehensive association studies continue to explore the intricate genetic interactions involving PRODH in psychiatric traits."}, {"type": "fg", "children": [{"type": "fg_f", "ref": "23"}]}, {"type": "t", "text": "\n"}]}, {"type": "t", "text": "\n\n"}, {"type": "p", "children": [{"type": "t", "text": "\nRecent work has also expanded the functional spectrum of PRODH beyond oncogenesis and neurodevelopment. Under conditions of metabolic stress, such as glucose deprivation or AMPK activation, enhanced PRODH activity facilitates the proline cycle to bolster ATP production and maintain redox equilibrium. In the cardiovascular context, diminished PRODH levels in failing hearts can be rescued by exercise, which in turn restores mitochondrial function and supports cardiomyocyte energy balance. These findings highlight PRODH as a versatile metabolic regulator that integrates nutrient availability with cell survival mechanisms across different tissues."}, {"type": "fg", "children": [{"type": "fg_f", "ref": "25"}]}, {"type": "t", "text": "\n"}]}, {"type": "rg", "children": [{"type": "r", "ref": 1, "children": [{"type": "t", "text": "Orianne Olivares, Jared R Mayers, Victoire Gouirand, et al. 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