{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,3]],"date-time":"2026-09-03T14:02:09Z","timestamp":1788444129037,"version":"build-2803163510"},"reference-count":58,"publisher":"Oxford University Press (OUP)","issue":"2","license":[{"start":{"date-parts":[[2017,9,23]],"date-time":"2017-09-23T00:00:00Z","timestamp":1506124800000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/academic.oup.com\/journals\/pages\/open_access\/funder_policies\/chorus\/standard_publication_model"}],"funder":[{"DOI":"10.13039\/100000069","name":"National Institute of Arthritis and Musculoskeletal and Skin Diseases","doi-asserted-by":"publisher","id":[{"id":"10.13039\/100000069","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2018,2,1]]},"abstract":"<jats:title>ABSTRACT<\/jats:title>\n                  <jats:sec>\n                    <jats:title>\u2003<\/jats:title>\n                    <jats:p>Intervertebral disc degeneration is a ubiquitous condition closely linked to chronic low-back pain. The health of the avascular nucleus pulposus (NP) plays a crucial role in the development of this pathology. We tested the hypothesis that a network comprising HIF-1\u03b1, carbonic anhydrase (CA) 9 and 12 isoforms, and sodium-coupled bicarbonate cotransporters (NBCs) buffer intracellular pH through coordinated bicarbonate recycling. Contrary to the current understanding of NP cell metabolism, analysis of metabolic-flux data from Seahorse XF analyzer showed that CO2 hydration contributes a significant source of extracellular proton production in NP cells, with a smaller input from glycolysis. Because enzymatic hydration of CO2 is catalyzed by plasma membrane-associated CAs we measured their expression and function in NP tissue. NP cells robustly expressed isoforms CA9\/12, which were hypoxia-inducible. In addition to increased mRNA stability under hypoxia, we observed binding of HIF-1\u03b1 to select hypoxia-responsive elements on CA9\/12 promoters using genomic chromatin immunoprecipitation. Importantly, in vitro loss of function studies and analysis of discs from NP-specific HIF-1\u03b1 null mice confirmed the dependency of CA9\/12 expression on HIF-1\u03b1. As expected, inhibition of CA activity decreased extracellular acidification rate independent of changes in HIF activity or lactate\/H+ efflux. Surprisingly, CA inhibition resulted in a concomitant decrease in intracellular pH that was mirrored by inhibition of sodium-bicarbonate importers. These results suggested that extracellular bicarbonate generated by CA9\/12 is recycled to buffer cytosolic pH fluctuations. Importantly, long-term intracellular acidification from CA inhibition lead to compromised cell viability, suggesting that plasma-membrane proton extrusion pathways alone are not sufficient to maintain homeostatic pH in NP cells. Taken together, our studies show for the first time that bicarbonate buffering through the HIF-1\u03b1\u2013CA axis is critical for NP cell survival in the hypoxic niche of the intervertebral disc. \u00a9 2017 American Society for Bone and Mineral Research.<\/jats:p>\n                  <\/jats:sec>","DOI":"10.1002\/jbmr.3293","type":"journal-article","created":{"date-parts":[[2017,9,23]],"date-time":"2017-09-23T03:03:18Z","timestamp":1506135798000},"page":"338-355","source":"Crossref","is-referenced-by-count":69,"title":["Bicarbonate Recycling by HIF-1\u2013Dependent Carbonic Anhydrase Isoforms 9 and 12 Is Critical in Maintaining Intracellular pH and Viability of Nucleus Pulposus Cells"],"prefix":"10.1093","volume":"33","author":[{"given":"Elizabeth S","family":"Silagi","sequence":"first","affiliation":[{"name":"Department of Orthopaedic Surgery, Sidney Kimmel Medical CollegeThomas Jefferson UniversityPhiladelphiaPAUSA"},{"name":"Program in Cell Biology and Regenerative Medicine, Jefferson College of Biomedical SciencesThomas Jefferson UniversityPhiladelphiaPAUSA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Zachary R","family":"Schoepflin","sequence":"additional","affiliation":[{"name":"Department of Orthopaedic Surgery, Sidney Kimmel Medical CollegeThomas Jefferson UniversityPhiladelphiaPAUSA"},{"name":"Program in Cell Biology and Regenerative Medicine, Jefferson College of Biomedical SciencesThomas Jefferson UniversityPhiladelphiaPAUSA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Erin L","family":"Seifert","sequence":"additional","affiliation":[{"name":"MitoCare Center, Department of Pathology, Anatomy and Cell BiologyThomas Jefferson UniversityPhiladelphiaPAUSA"},{"name":"Program in Cell Biology and Regenerative Medicine, Jefferson College of Biomedical SciencesThomas Jefferson UniversityPhiladelphiaPAUSA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Christophe","family":"Merceron","sequence":"additional","affiliation":[{"name":"Department of Orthopaedic SurgeryUniversity of Michigan Medical SchoolAnn ArborMIUSA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ernestina","family":"Schipani","sequence":"additional","affiliation":[{"name":"Department of MedicineDivision of EndocrinologyUniversity of Michigan Medical SchoolAnn ArborMIUSA"},{"name":"Department of Orthopaedic SurgeryUniversity of Michigan Medical SchoolAnn ArborMIUSA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Irving M","family":"Shapiro","sequence":"additional","affiliation":[{"name":"Department of Orthopaedic Surgery, Sidney Kimmel Medical CollegeThomas Jefferson UniversityPhiladelphiaPAUSA"},{"name":"Program in Cell Biology and Regenerative Medicine, Jefferson College of Biomedical SciencesThomas Jefferson UniversityPhiladelphiaPAUSA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Makarand V","family":"Risbud","sequence":"additional","affiliation":[{"name":"Department of Orthopaedic Surgery, Sidney Kimmel Medical CollegeThomas Jefferson UniversityPhiladelphiaPAUSA"},{"name":"Program in Cell Biology and Regenerative Medicine, Jefferson College of Biomedical SciencesThomas Jefferson UniversityPhiladelphiaPAUSA"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"286","published-online":{"date-parts":[[2017,9,23]]},"reference":[{"issue":"Suppl 2","key":"2024021014395108600_jbmr3293-bib-0001","first-page":"21","article-title":"Lumbar disc disorders and low-back pain: socioeconomic factors and consequences","volume":"88","author":"Katz","year":"2006","journal-title":"J Bone Joint Surg Am"},{"key":"2024021014395108600_jbmr3293-bib-0002","doi-asserted-by":"crossref","first-page":"591","DOI":"10.1001\/jama.2013.13805","article-title":"The state of US Health, 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