{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,11,10]],"date-time":"2026-11-10T09:54:22Z","timestamp":1794304462670,"version":"4.9.0"},"reference-count":84,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2025,12,12]],"date-time":"2025-12-12T00:00:00Z","timestamp":1765497600000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"},{"start":{"date-parts":[[2026,1,9]],"date-time":"2026-01-09T00:00:00Z","timestamp":1767916800000},"content-version":"vor","delay-in-days":28,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"}],"funder":[{"name":"National science foundation","award":["OCE-2049406"],"award-info":[{"award-number":["OCE-2049406"]}]},{"name":"National science foundation","award":["OCE-2049406"],"award-info":[{"award-number":["OCE-2049406"]}]},{"name":"National science foundation","award":["OCE-2049406"],"award-info":[{"award-number":["OCE-2049406"]}]}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Sci Rep"],"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>\n                    Coral reefs are essential for the foundation of marine ecosystems. However, ocean acidification (OA), driven by rising atmospheric carbon dioxide (CO\n                    <jats:sub>2<\/jats:sub>\n                    ) threatens coral growth and biological homeostasis. This study examines two Hawaiian coral species\u2014\n                    <jats:italic>Montipora capitata<\/jats:italic>\n                    and\n                    <jats:italic>Pocillopora acuta<\/jats:italic>\n                    to elevated pCO\n                    <jats:sub>2<\/jats:sub>\n                    simulating OA. Utilizing pH and O\n                    <jats:sub>2<\/jats:sub>\n                    microsensors under controlled light and dark conditions, this work characterized interspecific concentration boundary layer (CBL) traits and quantified material fluxes under ambient and elevated pCO\n                    <jats:sub>2<\/jats:sub>\n                    . The results of this study revealed that under increased pCO\n                    <jats:sub>2<\/jats:sub>\n                    ,\n                    <jats:italic>P. acuta<\/jats:italic>\n                    showed a significant reduction in dark proton efflux, followed by an increase in light O\n                    <jats:sub>2<\/jats:sub>\n                    flux, suggesting reduced calcification and enhanced photosynthesis. In contrast,\n                    <jats:italic>M. capitata<\/jats:italic>\n                    did not show any robust evidence of changes in either flux parameters under similar increased pCO\n                    <jats:sub>2<\/jats:sub>\n                    conditions. Statistical analyses using linear models revealed several significant interactions among species, treatment, and light conditions, identifying physical, chemical, and biological drivers of species responses to increased pCO\n                    <jats:sub>2<\/jats:sub>\n                    . This study also presents several conceptual models that correlate the CBL dynamics measured here with calcification and metabolic processes, thereby justifying our findings. We indicate that elevated pCO\n                    <jats:sub>2<\/jats:sub>\n                    exacerbates microchemical gradients in the CBL and may threaten calcification in vulnerable species such as\n                    <jats:italic>P. acuta<\/jats:italic>\n                    , while highlighting the resistance of\n                    <jats:italic>M. capitata<\/jats:italic>\n                    . Therefore, this study advances our understanding of how interspecific microenvironmental processes could influence coral responses to changing ocean chemistry.\n                  <\/jats:p>","DOI":"10.1038\/s41598-025-30818-4","type":"journal-article","created":{"date-parts":[[2025,12,12]],"date-time":"2025-12-12T15:36:08Z","timestamp":1765553768000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":2,"title":["Ocean acidification modulates material flux linked with coral calcification and photosynthesis"],"prefix":"10.1038","volume":"16","author":[{"given":"David A.","family":"Armstrong","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Conall","family":"McNicholl","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Keisha D.","family":"Bahr","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2025,12,12]]},"reference":[{"key":"30818_CR1","doi-asserted-by":"publisher","first-page":"365","DOI":"10.1038\/425365a","volume":"425","author":"K Caldeira","year":"2003","unstructured":"Caldeira, K. & Wickett, M. E. Anthropogenic carbon and ocean pH. Nature 425, 365\u2013365 (2003).","journal-title":"Nature"},{"key":"30818_CR2","doi-asserted-by":"publisher","first-page":"118","DOI":"10.1126\/science.284.5411.118","volume":"284","author":"JA Kleypas","year":"1999","unstructured":"Kleypas, J. A. et al. Geochemical consequences of increased atmospheric carbon dioxide on coral reefs. Science 284, 118\u2013120 (1999).","journal-title":"Science"},{"key":"30818_CR3","doi-asserted-by":"publisher","first-page":"367","DOI":"10.1126\/science.1097403","volume":"305","author":"CL Sabine","year":"2004","unstructured":"Sabine, C. L. et al. The oceanic sink for anthropogenic CO2. Science 305, 367\u2013371 (2004).","journal-title":"Science"},{"key":"30818_CR4","doi-asserted-by":"publisher","unstructured":"Changing & Ocean Marine Ecosystems, and Dependent Communities. in The Ocean and Cryosphere in a Changing Climate: Special Report of the Intergovernmental Panel on Climate Change (ed. Intergovernmental Panel on Climate Change (IPCC)) 447\u2013588Cambridge University Press, Cambridge, (2022). https:\/\/doi.org\/10.1017\/9781009157964.007","DOI":"10.1017\/9781009157964.007"},{"key":"30818_CR5","doi-asserted-by":"publisher","first-page":"414","DOI":"10.1093\/icesjms\/fsn048","volume":"65","author":"VJ Fabry","year":"2008","unstructured":"Fabry, V. J., Seibel, B. A., Feely, R. A. & Orr, J. C. Impacts of ocean acidification on marine fauna and ecosystem processes. ICES J. Mar. Sci. 65, 414\u2013432 (2008).","journal-title":"ICES J. Mar. Sci."},{"key":"30818_CR6","doi-asserted-by":"publisher","first-page":"160","DOI":"10.1093\/icb\/39.1.160","volume":"39","author":"JP Gattuso","year":"1999","unstructured":"Gattuso, J. P., Allemand, D. & Frankignoulle, M. Photosynthesis and calcification at Cellular, organismal and community levels in coral reefs: A review on interactions and control by carbonate chemistry. Am. Zool. 39, 160\u2013183 (1999).","journal-title":"Am. Zool."},{"key":"30818_CR7","doi-asserted-by":"publisher","first-page":"365","DOI":"10.5194\/bg-12-365-2015","volume":"12","author":"S Comeau","year":"2015","unstructured":"Comeau, S., Carpenter, R. C., Lantz, C. A. & Edmunds, P. J. Ocean acidification accelerates dissolution of experimental coral reef communities. Biogeosciences 12, 365\u2013372 (2015).","journal-title":"Biogeosciences"},{"key":"30818_CR8","doi-asserted-by":"publisher","first-page":"745","DOI":"10.1002\/2016JC012326","volume":"122","author":"TM DeCarlo","year":"2017","unstructured":"DeCarlo, T. M. et al. Community production modulates coral reef pH and the sensitivity of ecosystem calcification to ocean acidification. J. Geophys. Res. Oceans. 122, 745\u2013761 (2017).","journal-title":"J. Geophys. Res. Oceans"},{"key":"30818_CR9","doi-asserted-by":"publisher","first-page":"e378","DOI":"10.7717\/peerj.378","volume":"2","author":"PL Jokiel","year":"2014","unstructured":"Jokiel, P. L., Jury, C. P. & Rodgers, K. S. Coral-algae metabolism and diurnal changes in the CO2-carbonate system of bulk sea water. PeerJ 2, e378 (2014).","journal-title":"PeerJ"},{"key":"30818_CR10","doi-asserted-by":"publisher","first-page":"321","DOI":"10.1146\/annurev-marine-121211-172241","volume":"5","author":"AJ Andersson","year":"2013","unstructured":"Andersson, A. J. & Gledhill, D. Ocean acidification and coral reefs: effects on Breakdown, Dissolution, and net ecosystem calcification. Annu. Rev. Mar. Sci. 5, 321\u2013348 (2013).","journal-title":"Annu. Rev. Mar. Sci."},{"key":"30818_CR11","doi-asserted-by":"publisher","first-page":"473","DOI":"10.1007\/s00338-008-0380-9","volume":"27","author":"PL Jokiel","year":"2008","unstructured":"Jokiel, P. L. et al. Ocean acidification and calcifying reef organisms: a mesocosm investigation. Coral Reefs. 27, 473\u2013483 (2008).","journal-title":"Coral Reefs"},{"key":"30818_CR12","doi-asserted-by":"publisher","first-page":"729","DOI":"10.1007\/s00338-016-1405-4","volume":"35","author":"KD Bahr","year":"2016","unstructured":"Bahr, K. D., Jokiel, P. L. & Rodgers, K. S. Relative sensitivity of five Hawaiian coral species to high temperature under high-pCO2 conditions. Coral Reefs. 35, 729\u2013738 (2016).","journal-title":"Coral Reefs"},{"key":"30818_CR13","doi-asserted-by":"publisher","first-page":"1083","DOI":"10.1093\/icesjms\/fsw078","volume":"74","author":"KD Bahr","year":"2017","unstructured":"Bahr, K. D., Jokiel, P. L. & Rodgers, K. S. Seasonal and annual calcification rates of the Hawaiian reef coral, Montipora capitata, under present and future climate change scenarios. ICES J. Mar. Sci. 74, 1083\u20131091 (2017).","journal-title":"ICES J. Mar. Sci."},{"key":"30818_CR14","doi-asserted-by":"publisher","first-page":"20182840","DOI":"10.1098\/rspb.2018.2840","volume":"286","author":"CB Bove","year":"2019","unstructured":"Bove, C. B. et al. Common Caribbean corals exhibit highly variable responses to future acidification and warming. Proc. R. Soc. B Biol. Sci. 286, 20182840 (2019).","journal-title":"Proc. R. Soc. B Biol. Sci."},{"key":"30818_CR15","doi-asserted-by":"publisher","first-page":"477","DOI":"10.1038\/s41558-019-0486-9","volume":"9","author":"S Comeau","year":"2019","unstructured":"Comeau, S. et al. Resistance to ocean acidification in coral reef taxa is not gained by acclimatization. Nat. Clim. Change. 9, 477\u2013483 (2019).","journal-title":"Nat. Clim. Change"},{"key":"30818_CR16","doi-asserted-by":"publisher","first-page":"388","DOI":"10.4319\/lo.2013.58.1.0388","volume":"58","author":"S Comeau","year":"2013","unstructured":"Comeau, S., Edmunds, P. J., Spindel, N. B. & Carpenter, R. C. The responses of eight coral reef calcifiers to increasing partial pressure of CO2 do not exhibit a tipping point. Limnol. Oceanogr. 58, 388\u2013398 (2013).","journal-title":"Limnol. Oceanogr."},{"key":"30818_CR17","doi-asserted-by":"publisher","first-page":"924","DOI":"10.1080\/17451000.2018.1551616","volume":"14","author":"KD Bahr","year":"2018","unstructured":"Bahr, K. D., Rodgers, K. S. & Jokiel, P. L. Ocean warming drives decline in coral metabolism while acidification highlights species-specific responses. Mar. Biol. Res. 14, 924\u2013935 (2018).","journal-title":"Mar. Biol. Res."},{"key":"30818_CR18","doi-asserted-by":"publisher","first-page":"12829","DOI":"10.1038\/s41598-019-49044-w","volume":"9","author":"S Comeau","year":"2019","unstructured":"Comeau, S. et al. Flow-driven micro-scale pH variability affects the physiology of corals and coralline algae under ocean acidification. Sci. Rep. 9, 12829 (2019).","journal-title":"Sci. Rep."},{"key":"30818_CR19","doi-asserted-by":"crossref","unstructured":"Chan, N. C. S., Wangpraseurt, D., K\u00fchl, M. & Connolly, S. R. Flow and coral morphology control coral surface pH: implications for the effects of ocean acidification. Front Mar. Sci 3, 10 (2016).","DOI":"10.3389\/fmars.2016.00010"},{"key":"30818_CR20","doi-asserted-by":"publisher","first-page":"282","DOI":"10.1111\/gcb.12011","volume":"19","author":"NCS Chan","year":"2013","unstructured":"Chan, N. C. S. & Connolly, S. R. Sensitivity of coral calcification to ocean acidification: a meta-analysis. Glob Change Biol. 19, 282\u2013290 (2013).","journal-title":"Glob Change Biol."},{"key":"30818_CR21","doi-asserted-by":"publisher","first-page":"1660","DOI":"10.1046\/j.1365-2486.2003.00678.x","volume":"9","author":"S Reynaud","year":"2003","unstructured":"Reynaud, S. et al. Interacting effects of CO2 partial pressure and temperature on photosynthesis and calcification in a scleractinian coral. Glob Change Biol. 9, 1660\u20131668 (2003).","journal-title":"Glob Change Biol."},{"key":"30818_CR22","doi-asserted-by":"crossref","unstructured":"Langdon, C. & Atkinson, M. J. Effect of elevated pCO2 on photosynthesis and calcification of corals and interactions with seasonal change in temperature\/irradiance and nutrient enrichment. J Geophys. Res. Oceans 110, C09S07 (2005).","DOI":"10.1029\/2004JC002576"},{"key":"30818_CR23","doi-asserted-by":"publisher","first-page":"7","DOI":"10.1007\/978-94-017-7567-0_2","volume-title":"Coral Reefs at the Crossroads","author":"PL Jokiel","year":"2016","unstructured":"Jokiel, P. L., Jury, C. P. & Kuffner, I. B. Coral calcification and ocean acidification. In Coral Reefs at the Crossroads Vol. 6 (eds Hubbard, D. K. et al.) 7\u201345 (Springer Netherlands, 2016)."},{"key":"30818_CR24","doi-asserted-by":"publisher","first-page":"177","DOI":"10.1007\/978-94-007-0114-4_11","volume-title":"Coral Reefs: an Ecosystem in Transition","author":"JA Kaandorp","year":"2011","unstructured":"Kaandorp, J. A., Filatov, M. & Chindapol, N. Simulating and quantifying the environmental influence on coral colony growth and form. In Coral Reefs: an Ecosystem in Transition (eds Dubinsky, Z. & Stambler, N.) 177\u2013185 (Springer Netherlands, 2011). https:\/\/doi.org\/10.1007\/978-94-007-0114-4_11."},{"key":"30818_CR25","doi-asserted-by":"publisher","first-page":"12757","DOI":"10.1038\/s41598-024-63210-9","volume":"14","author":"CPP Martins","year":"2024","unstructured":"Martins, C. P. P., Ziegler, M., Schubert, P., Wilke, T. & Wall, M. Effects of water flow and ocean acidification on oxygen and pH gradients in coral boundary layer. Sci. Rep. 14, 12757 (2024).","journal-title":"Sci. Rep."},{"key":"30818_CR26","doi-asserted-by":"publisher","first-page":"639","DOI":"10.5343\/bms.2010.1107","volume":"87","author":"PL Jokiel","year":"2011","unstructured":"Jokiel, P. L. Ocean acidification and control of reef coral calcification by boundary layer limitation of proton flux. Bull. Mar. Sci. 87, 639\u2013657 (2011).","journal-title":"Bull. Mar. Sci."},{"key":"30818_CR27","doi-asserted-by":"publisher","first-page":"17","DOI":"10.1016\/0022-0981(95)00156-5","volume":"199","author":"N Shashar","year":"1996","unstructured":"Shashar, N., Kinane, S., Jokiel, P. L. & Patterson, M. R. Hydromechanical boundary layers over a coral reef. J. Exp. Mar. Biol. Ecol. 199, 17\u201328 (1996).","journal-title":"J. Exp. Mar. Biol. Ecol."},{"key":"30818_CR28","doi-asserted-by":"publisher","first-page":"e97235","DOI":"10.1371\/journal.pone.0097235","volume":"9","author":"CE Cornwall","year":"2014","unstructured":"Cornwall, C. E. et al. Diffusion boundary layers ameliorate the negative effects of ocean acidification on the temperate coralline macroalga arthrocardia corymbosa. PLOS ONE. 9, e97235 (2014).","journal-title":"PLOS ONE"},{"key":"30818_CR29","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1038\/s42003-024-05938-8","volume":"7","author":"L Crovetto","year":"2024","unstructured":"Crovetto, L., Venn, A. A., Sevilgen, D., Tambutt\u00e9, S. & Tambutt\u00e9, E. Spatial variability of and effect of light on the c\u0153lenteron pH of a reef coral. Commun. Biol. 7, 1\u201310 (2024).","journal-title":"Commun. Biol."},{"key":"30818_CR30","doi-asserted-by":"publisher","first-page":"1785","DOI":"10.1098\/rsif.2011.0144","volume":"8","author":"IM Jimenez","year":"2011","unstructured":"Jimenez, I. M., K\u00fchl, M., Larkum, A. W. D. & Ralph, P. J. Effects of flow and colony morphology on the thermal boundary layer of corals. J. R Soc. Interface. 8, 1785\u20131795 (2011).","journal-title":"J. R Soc. Interface"},{"key":"30818_CR31","doi-asserted-by":"publisher","DOI":"10.1007\/s00338-024-02531-7","author":"CPP Martins","year":"2024","unstructured":"Martins, C. P. P., Wall, M., Schubert, P., Wilke, T. & Ziegler, M. Variability of the surface boundary layer of reef-building coral species. Coral Reefs. https:\/\/doi.org\/10.1007\/s00338-024-02531-7 (2024).","journal-title":"Coral Reefs"},{"key":"30818_CR32","doi-asserted-by":"publisher","first-page":"4150","DOI":"10.1016\/j.cub.2022.07.071","volume":"32","author":"CO Pacherres","year":"2022","unstructured":"Pacherres, C. O., Ahmerkamp, S., Koren, K., Richter, C. & Holtappels, M. Ciliary flows in corals ventilate target areas of high photosynthetic oxygen production. Curr. Biol. 32, 4150\u20134158e3 (2022).","journal-title":"Curr. Biol."},{"key":"30818_CR33","doi-asserted-by":"publisher","first-page":"7541","DOI":"10.1038\/s41598-020-64420-7","volume":"10","author":"CO Pacherres","year":"2020","unstructured":"Pacherres, C. O., Ahmerkamp, S., Schmidt-Grieb, G. M., Holtappels, M. & Richter, C. Ciliary vortex flows and oxygen dynamics in the coral boundary layer. Sci. Rep. 10, 7541 (2020).","journal-title":"Sci. Rep."},{"key":"30818_CR34","doi-asserted-by":"publisher","first-page":"12525","DOI":"10.1038\/s41598-021-91030-8","volume":"11","author":"EM Henley","year":"2021","unstructured":"Henley, E. M. et al. Reproductive plasticity of Hawaiian Montipora corals following thermal stress. Sci. Rep. 11, 12525 (2021).","journal-title":"Sci. Rep."},{"key":"30818_CR35","doi-asserted-by":"publisher","first-page":"931","DOI":"10.1007\/s00338-023-02397-1","volume":"42","author":"J Saper","year":"2023","unstructured":"Saper, J., H\u00f8j, L., Humphrey, C. & Bourne, D. G. Quantifying capture and ingestion of live feeds across three coral species. Coral Reefs. 42, 931\u2013943 (2023).","journal-title":"Coral Reefs"},{"key":"30818_CR36","doi-asserted-by":"publisher","first-page":"1297483","DOI":"10.3389\/fgene.2023.1297483","volume":"14","author":"T Han","year":"2023","unstructured":"Han, T. et al. Comparative transcriptome analysis reveals deep molecular landscapes in stony coral Montipora clade. Front. Genet. 14, 1297483 (2023).","journal-title":"Front. Genet."},{"key":"30818_CR37","doi-asserted-by":"publisher","first-page":"1976","DOI":"10.1002\/lno.12181","volume":"67","author":"D Bhattacharya","year":"2022","unstructured":"Bhattacharya, D., Stephens, T. G., Tinoco, A. I., Richmond, R. H. & Cleves, P. A. Life on the edge: Hawaiian model for coral evolution. Limnol. Oceanogr. 67, 1976\u20131985 (2022).","journal-title":"Limnol. Oceanogr."},{"key":"30818_CR38","doi-asserted-by":"publisher","first-page":"1169","DOI":"10.1007\/s00338-018-1726-6","volume":"37","author":"V Schoepf","year":"2018","unstructured":"Schoepf, V. et al. Impacts of coral bleaching on pH and oxygen gradients across the coral concentration boundary layer: a microsensor study. Coral Reefs. 37, 1169\u20131180 (2018).","journal-title":"Coral Reefs"},{"key":"30818_CR39","doi-asserted-by":"publisher","first-page":"37","DOI":"10.1016\/S0921-8181(98)00035-6","volume":"18","author":"J Gattuso","year":"1998","unstructured":"Gattuso, J. Effect of calcium carbonate saturation of seawater on coral calcification. Glob Planet. Change. 18, 37\u201346 (1998).","journal-title":"Glob Planet. Change"},{"key":"30818_CR40","doi-asserted-by":"publisher","first-page":"639","DOI":"10.1029\/1999GB001195","volume":"14","author":"C Langdon","year":"2000","unstructured":"Langdon, C. et al. Effect of calcium carbonate saturation state on the calcification rate of an experimental coral reef. Glob Biogeochem. Cycles. 14, 639\u2013654 (2000).","journal-title":"Glob Biogeochem. Cycles"},{"key":"30818_CR41","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1016\/j.jembe.2011.10.008","volume":"409","author":"PL Jokiel","year":"2011","unstructured":"Jokiel, P. L. The reef coral two compartment proton flux model: A new approach relating tissue-level physiological processes to gross corallum morphology. J. Exp. Mar. Biol. Ecol. 409, 1\u201312 (2011).","journal-title":"J. Exp. Mar. Biol. Ecol."},{"key":"30818_CR42","doi-asserted-by":"publisher","first-page":"e17186","DOI":"10.1111\/mec.17186","volume":"33","author":"VZ Radice","year":"2024","unstructured":"Radice, V. Z., Martinez, A., Paytan, A., Potts, D. C. & Barshis, D. J. Complex dynamics of coral gene expression responses to low pH across species. Mol. Ecol. 33, e17186 (2024).","journal-title":"Mol. Ecol."},{"key":"30818_CR43","doi-asserted-by":"publisher","first-page":"151","DOI":"10.1007\/s10126-018-9864-y","volume":"21","author":"X Yuan","year":"2019","unstructured":"Yuan, X. et al. Gene expression profiles of two coral species with varied resistance to ocean acidification. Mar. Biotechnol. 21, 151\u2013160 (2019).","journal-title":"Mar. Biotechnol."},{"key":"30818_CR44","doi-asserted-by":"publisher","first-page":"1632","DOI":"10.1111\/j.1365-2486.2009.02057.x","volume":"16","author":"CP Jury","year":"2010","unstructured":"Jury, C. P., Whitehead, R. F. & Szmant, A. M. Effects of variations in carbonate chemistry on the calcification rates of Madracis Auretenra (=\u2009Madracis mirabilis sensu Wells, 1973): bicarbonate concentrations best predict calcification rates. Glob Change Biol. 16, 1632\u20131644 (2010).","journal-title":"Glob Change Biol."},{"key":"30818_CR45","doi-asserted-by":"publisher","DOI":"10.5194\/bgd-12-6689-2015","author":"LT Bach","year":"2015","unstructured":"Bach, L. T. Reconsidering the role of carbonate ion concentration in calcification by marine organisms. Preprint at. https:\/\/doi.org\/10.5194\/bgd-12-6689-2015 (2015).","journal-title":"Preprint at."},{"key":"30818_CR46","doi-asserted-by":"crossref","unstructured":"Hohn, S. & Merico, A. Quantifying the relative importance of transcellular and paracellular ion transports to coral polyp calcification. Front. Earth Sci. 2, 37 (2015).","DOI":"10.3389\/feart.2014.00037"},{"key":"30818_CR47","doi-asserted-by":"crossref","unstructured":"Venn, A. A., Tambutt\u00e9, E., Comeau, S. & Tambutt\u00e9, S. Proton gradients across the coral calcifying cell layer: effects of light, ocean acidification and carbonate chemistry. Front Mar. Sci 9, 973908 (2022).","DOI":"10.3389\/fmars.2022.973908"},{"key":"30818_CR48","doi-asserted-by":"publisher","first-page":"973908","DOI":"10.3389\/fmars.2022.973908","volume":"9","author":"AA Venn","year":"2022","unstructured":"Venn, A. A., Tambutt\u00e9, E., Comeau, S. & Tambutt\u00e9, S. Proton gradients across the coral calcifying cell layer: effects of light, ocean acidification and carbonate chemistry. Front. Mar. Sci. 9, 973908 (2022).","journal-title":"Front. Mar. Sci."},{"key":"30818_CR49","doi-asserted-by":"publisher","first-page":"1634","DOI":"10.1073\/pnas.1216153110","volume":"110","author":"AA Venn","year":"2013","unstructured":"Venn, A. A. et al. Impact of seawater acidification on pH at the tissue\u2013skeleton interface and calcification in reef corals. Proc. Natl. Acad. Sci. 110, 1634\u20131639 (2013).","journal-title":"Proc. Natl. Acad. Sci."},{"key":"30818_CR50","doi-asserted-by":"publisher","first-page":"193","DOI":"10.3390\/oceans2010012","volume":"2","author":"CE Reymond","year":"2021","unstructured":"Reymond, C. E. & Hohn, S. An experimental approach to assessing the roles of Magnesium, Calcium, and carbonate ratios in marine carbonates. Oceans 2, 193\u2013214 (2021).","journal-title":"Oceans"},{"key":"30818_CR51","doi-asserted-by":"publisher","first-page":"1092","DOI":"10.1093\/icesjms\/fsv267","volume":"74","author":"S Comeau","year":"2017","unstructured":"Comeau, S., Carpenter, R. C. & Edmunds, P. J. Effects of pCO2 on photosynthesis and respiration of tropical scleractinian corals and calcified algae. ICES J. Mar. Sci. 74, 1092\u20131102 (2017).","journal-title":"ICES J. Mar. Sci."},{"key":"30818_CR52","doi-asserted-by":"publisher","first-page":"e34659","DOI":"10.1371\/journal.pone.0034659","volume":"7","author":"P Kaniewska","year":"2012","unstructured":"Kaniewska, P. et al. Major cellular and physiological impacts of ocean acidification on a reef Building coral. PLoS ONE. 7, e34659 (2012).","journal-title":"PLoS ONE"},{"key":"30818_CR53","doi-asserted-by":"publisher","first-page":"179","DOI":"10.1016\/j.cbpa.2015.02.018","volume":"184","author":"J Strahl","year":"2015","unstructured":"Strahl, J. et al. Physiological and ecological performance differs in four coral taxa at a volcanic carbon dioxide seep. Comp. Biochem. Physiol. Mol. Integr. Physiol. 184, 179\u2013186 (2015).","journal-title":"Comp. Biochem. Physiol. Mol. Integr. Physiol."},{"key":"30818_CR54","doi-asserted-by":"publisher","first-page":"6681","DOI":"10.1038\/srep06681","volume":"4","author":"S Comeau","year":"2014","unstructured":"Comeau, S., Edmunds, P. J., Lantz, C. A. & Carpenter, R. C. Water flow modulates the response of coral reef communities to ocean acidification. Sci. Rep. 4, 6681 (2014).","journal-title":"Sci. Rep."},{"key":"30818_CR55","doi-asserted-by":"publisher","first-page":"7573","DOI":"10.1038\/s41598-017-08003-z","volume":"7","author":"S Comeau","year":"2017","unstructured":"Comeau, S., Cornwall, C. E. & McCulloch, M. T. Decoupling between the response of coral calcifying fluid pH and calcification to ocean acidification. Sci. Rep. 7, 7573 (2017).","journal-title":"Sci. Rep."},{"key":"30818_CR56","doi-asserted-by":"publisher","first-page":"20181168","DOI":"10.1098\/rspb.2018.1168","volume":"285","author":"CE Cornwall","year":"2018","unstructured":"Cornwall, C. E. et al. Resistance of corals and coralline algae to ocean acidification: physiological control of calcification under natural pH variability. Proc. R. Soc. B Biol. Sci. 285, 20181168 (2018).","journal-title":"Proc. R. Soc. B Biol. Sci."},{"key":"30818_CR57","doi-asserted-by":"publisher","first-page":"4857","DOI":"10.1111\/gcb.14379","volume":"24","author":"S Comeau","year":"2018","unstructured":"Comeau, S., Cornwall, C. E., DeCarlo, T. M., Krieger, E. & McCulloch, M. T. Similar controls on calcification under ocean acidification across unrelated coral reef taxa. Glob Change Biol. 24, 4857\u20134868 (2018).","journal-title":"Glob Change Biol."},{"key":"30818_CR58","doi-asserted-by":"publisher","first-page":"453","DOI":"10.1016\/j.crpv.2004.07.011","volume":"3","author":"D Allemand","year":"2004","unstructured":"Allemand, D. et al. Biomineralisation in reef-building corals: from molecular mechanisms to environmental control. C.R. Palevol. 3, 453\u2013467 (2004).","journal-title":"C.R. Palevol"},{"key":"30818_CR59","doi-asserted-by":"publisher","first-page":"95","DOI":"10.1016\/S0012-8252(96)00036-0","volume":"42","author":"TA McConnaughey","year":"1997","unstructured":"McConnaughey, T. A. & Whelan, J. F. Calcification generates protons for nutrient and bicarbonate uptake. Earth-Sci. Rev. 42, 95\u2013117 (1997).","journal-title":"Earth-Sci. Rev."},{"key":"30818_CR60","doi-asserted-by":"publisher","first-page":"1437","DOI":"10.1016\/j.bmc.2012.10.024","volume":"21","author":"A Bertucci","year":"2013","unstructured":"Bertucci, A. et al. Carbonic anhydrases in anthozoan corals\u2014A review. Bioorg. Med. Chem. 21, 1437\u20131450 (2013).","journal-title":"Bioorg. Med. Chem."},{"key":"30818_CR61","doi-asserted-by":"publisher","first-page":"119","DOI":"10.1007\/978-94-007-0114-4_9","volume-title":"Coral Reefs: an Ecosystem in Transition","author":"D Allemand","year":"2011","unstructured":"Allemand, D., Tambutt\u00e9, \u00c9., Zoccola, D. & Tambutt\u00e9, S. Coral Calcification, cells to reefs. In Coral Reefs: an Ecosystem in Transition (eds Dubinsky, Z. & Stambler, N.) 119\u2013150 (Springer Netherlands, 2011). https:\/\/doi.org\/10.1007\/978-94-007-0114-4_9."},{"key":"30818_CR62","doi-asserted-by":"publisher","first-page":"3445","DOI":"10.1242\/jeb.203.22.3445","volume":"203","author":"P Furla","year":"2000","unstructured":"Furla, P., Galgani, I., Durand, I. & Allemand, D. Sources and mechanisms of inorganic carbon transport for coral calcification and photosynthesis. J. Exp. Biol. 203, 3445\u20133457 (2000).","journal-title":"J. Exp. Biol."},{"key":"30818_CR63","first-page":"925","volume":"76","author":"D Allemand","year":"1998","unstructured":"Allemand, D., Furla, P. & B\u00e9nazet-Tambutt\u00e9, S. Mechanisms of carbon acquisition for endosymbiont photosynthesis in anthozoa. Can. J. Bot. 76, 925\u2013941 (1998).","journal-title":"Can. J. Bot."},{"key":"30818_CR64","doi-asserted-by":"publisher","first-page":"1106","DOI":"10.3390\/jmse10081106","volume":"10","author":"LP Cameron","year":"2022","unstructured":"Cameron, L. P. et al. Impacts of warming and acidification on coral calcification linked to photosymbiont loss and deregulation of calcifying fluid pH. J. Mar. Sci. Eng. 10, 1106 (2022).","journal-title":"J. Mar. Sci. Eng."},{"key":"30818_CR65","doi-asserted-by":"publisher","first-page":"607","DOI":"10.1073\/pnas.1413483112","volume":"112","author":"KL Barott","year":"2015","unstructured":"Barott, K. L, Venn, A. A, Perez, S. O., Tambutt\u00e9S S. & Tresguerres M. Coral host cells acidify symbiotic algal microenvironment to promote photosynthesis. Proc. Natl. Acad. Sci. 112, 607\u2013612 (2015).","journal-title":"Proc. Natl. Acad. Sci."},{"key":"30818_CR66","doi-asserted-by":"publisher","first-page":"271","DOI":"10.1007\/s40641-017-0082-x","volume":"3","author":"EB Rivest","year":"2017","unstructured":"Rivest, E. B., Comeau, S. & Cornwall, C. E. The role of natural variability in shaping the response of coral reef organisms to climate change. Curr. Clim. Change Rep. 3, 271\u2013281 (2017).","journal-title":"Curr. Clim. Change Rep."},{"key":"30818_CR67","doi-asserted-by":"publisher","first-page":"1632","DOI":"10.1111\/gcb.12154","volume":"19","author":"EC Shaw","year":"2013","unstructured":"Shaw, E. C., McNeil, B. I., Tilbrook, B., Matear, R. & Bates, M. L. Anthropogenic changes to seawater buffer capacity combined with natural reef metabolism induce extreme future coral reef CO2 conditions. Glob Change Biol. 19, 1632\u20131641 (2013).","journal-title":"Glob Change Biol."},{"key":"30818_CR68","doi-asserted-by":"publisher","first-page":"135","DOI":"10.1016\/j.jembe.2018.06.007","volume":"506","author":"IC Enochs","year":"2018","unstructured":"Enochs, I. C. et al. The influence of diel carbonate chemistry fluctuations on the calcification rate of Acropora cervicornis under present day and future acidification conditions. J. Exp. Mar. Biol. Ecol. 506, 135\u2013143 (2018).","journal-title":"J. Exp. Mar. Biol. Ecol."},{"key":"30818_CR69","doi-asserted-by":"publisher","first-page":"155","DOI":"10.1007\/BF00265006","volume":"8","author":"PL Jokiel","year":"1990","unstructured":"Jokiel, P. L. & Coles, S. L. Response of Hawaiian and other Indo-Pacific reef corals to elevated temperature. Coral Reefs. 8, 155\u2013162 (1990).","journal-title":"Coral Reefs"},{"key":"30818_CR70","doi-asserted-by":"publisher","first-page":"499","DOI":"10.1002\/2013GL058489","volume":"41","author":"KEF Shamberger","year":"2014","unstructured":"Shamberger, K. E. F. et al. Diverse coral communities in naturally acidified waters of a Western Pacific reef. Geophys. Res. Lett. 41, 499\u2013504 (2014).","journal-title":"Geophys. Res. Lett."},{"key":"30818_CR71","doi-asserted-by":"publisher","first-page":"99","DOI":"10.3354\/meps10690","volume":"501","author":"S Comeau","year":"2014","unstructured":"Comeau, S., Edmunds, P. J., Spindel, N. B. & Carpenter, R. C. Diel pCO2 oscillations modulate the response of the coral acropora hyacinthus to ocean acidification. Mar. Ecol. Prog Ser. 501, 99\u2013111 (2014).","journal-title":"Mar. Ecol. Prog Ser."},{"key":"30818_CR72","doi-asserted-by":"publisher","first-page":"9851","DOI":"10.1029\/2019GL083726","volume":"46","author":"AK Kealoha","year":"2019","unstructured":"Kealoha, A. K. et al. Heterotrophy of oceanic particulate organic matter elevates net ecosystem calcification. Geophys. Res. Lett. 46, 9851\u20139860 (2019).","journal-title":"Geophys. Res. Lett."},{"key":"30818_CR73","doi-asserted-by":"publisher","first-page":"e950","DOI":"10.7717\/peerj.950","volume":"3","author":"KD Bahr","year":"2015","unstructured":"Bahr, K. D., Jokiel, P. L. & Toonen, R. J. The unnatural history of K\u0101ne\u2018ohe bay: coral reef resilience in the face of centuries of anthropogenic impacts. PeerJ 3, e950 (2015).","journal-title":"PeerJ"},{"key":"30818_CR74","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1111\/zoj.12092","volume":"170","author":"S Schmidt-Roach","year":"2014","unstructured":"Schmidt-Roach, S., Miller, K. J., Lundgren, P. & Andreakis, N. With eyes wide open: a revision of species within and closely related to the ocillopora damicornis species complex (Scleractinia; Pocilloporidae) using morphology and genetics. Zool. J. Linn. Soc. 170, 1\u201333 (2014).","journal-title":"Zool. J. Linn. Soc."},{"key":"30818_CR75","doi-asserted-by":"publisher","first-page":"262","DOI":"10.1016\/j.zool.2013.06.001","volume":"116","author":"DM Yost","year":"2013","unstructured":"Yost, D. M. et al. Diversity in skeletal architecture influences biological heterogeneity and Symbiodinium habitat in corals. Zoology 116, 262\u2013269 (2013).","journal-title":"Zoology"},{"key":"30818_CR76","doi-asserted-by":"crossref","unstructured":"Cohen, A. L., McCorkle, D. C., de Putron, S., Gaetani, G. A. & Rose, K. A. Morphological and compositional changes in the skeletons of new coral recruits reared in acidified seawater: insights into the biomineralization response to ocean acidification. Geochem Geophys. Geosystems 10, Q07005 (2009).","DOI":"10.1029\/2009GC002411"},{"key":"30818_CR77","doi-asserted-by":"crossref","unstructured":"Shapiro, O. et al. Vortical ciliary flows actively enhance mass transport in reef corals. Proc. Natl. Acad. Sci. 111, (2014).","DOI":"10.1073\/pnas.1323094111"},{"key":"30818_CR78","doi-asserted-by":"publisher","first-page":"455","DOI":"10.2307\/1542485","volume":"185","author":"N Shashar","year":"1993","unstructured":"Shashar, N., Cohen, Y. & Loya, Y. Extreme diel fluctuations of oxygen in diffusive boundary layers surrounding stony corals. Biol. Bull. 185, 455\u2013461 (1993).","journal-title":"Biol. Bull."},{"key":"30818_CR79","doi-asserted-by":"publisher","first-page":"159","DOI":"10.3354\/meps117159","volume":"117","author":"M K\u00fchl","year":"1995","unstructured":"K\u00fchl, M., Cohen, Y., Dalsgaard, T., J\u00f8rgensen, B. B. & Revsbech, N. P. Microenvironment and photosynthesis of zooxanthellae in scleractinian corals studied with microsensors for O\u2082, pH and light. Mar. Ecol. Prog Ser. 117, 159\u2013172 (1995).","journal-title":"Mar. Ecol. Prog Ser."},{"key":"30818_CR80","unstructured":"Jokiel, P. Coral growth: buoyant weight technique. Coral Reefs: Research Methods 529\u2013541 (1978)."},{"key":"30818_CR81","doi-asserted-by":"publisher","first-page":"313","DOI":"10.4319\/lom.2014.12.313","volume":"12","author":"PL Jokiel","year":"2014","unstructured":"Jokiel, P. L., Bahr, K. D. & Rodgers, K. S. Low-cost, high-flow mesocosm system for simulating ocean acidification with CO2 gas. Limnol. Oceanogr. Methods. 12, 313\u2013322 (2014).","journal-title":"Limnol. Oceanogr. Methods"},{"key":"30818_CR82","unstructured":"Guide to Best Practices for Ocean CO2 Measurements (North Pacific Marine Science Organization, 2007)."},{"key":"30818_CR83","doi-asserted-by":"publisher","first-page":"341","DOI":"10.1111\/j.1529-8817.2011.00958.x","volume":"47","author":"CL Hurd","year":"2011","unstructured":"Hurd, C. L. & Pilditch, C. A. Flow-Induced morphological variations affect diffusion Boundary-Layer thickness of macrocystis pyrifera (heterokontophyta, Laminariales)1. J. Phycol. 47, 341\u2013351 (2011).","journal-title":"J. Phycol."},{"key":"30818_CR84","unstructured":"R Core Team. The R Project for Statistical Computing. https:\/\/www.r-project.org\/"}],"container-title":["Scientific Reports"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.nature.com\/articles\/s41598-025-30818-4.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41598-025-30818-4","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41598-025-30818-4.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,1,11]],"date-time":"2026-01-11T21:04:04Z","timestamp":1768165444000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.nature.com\/articles\/s41598-025-30818-4"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2025,12,12]]},"references-count":84,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2026,12]]}},"alternative-id":["30818"],"URL":"https:\/\/doi.org\/10.1038\/s41598-025-30818-4","relation":{"has-preprint":[{"id-type":"doi","id":"10.21203\/rs.3.rs-6149474\/v1","asserted-by":"object"}]},"ISSN":["2045-2322"],"issn-type":[{"value":"2045-2322","type":"electronic"}],"subject":[],"published":{"date-parts":[[2025,12,12]]},"assertion":[{"value":"3 March 2025","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"27 November 2025","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"12 December 2025","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"order":1,"name":"Ethics","group":{"name":"EthicsHeading","label":"Declarations"}},{"value":"The authors declare no competing interests.","order":2,"name":"Ethics","group":{"name":"EthicsHeading","label":"Competing interests"}}],"article-number":"1255"}}