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30 , 31 Further studies demonstrated that in addition to the hypoxic induction of EPO in the kidney and liver, HIF-1 played a more general role, regulating hypoxia-sensitive genes, including those encoding glycolytic enzymes (e.g., phosphoglycerate kinase 1 and lactate dehydrogenase A) and angiogenic factors, such as vascular endothelial growth factor (VEGF).32, 33, 34, 35 Subsequently, HIF transcriptional activity was found to be controlled by hydroxylation of specific proline residues within the oxygen-regulated HIF-1α subunit, leading to its ubiquitylation and proteasomal degradation.36, 37, 38 HIF-α hydroxylation is mediated by a family of prolyl-4-hydroxylases, PHD enzymes, which function as the primary oxygen sensors of the HIF pathway.39, 40, 41, 42 In 2019, the Nobel Prize in Physiology or Medicine was awarded to Professor William Kaelin, Jr., Sir Peter Ratcliffe, and Gregg Semenza for their contributions to delineating the molecular mechanisms underlying HIF oxygen sensing
