{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,16]],"date-time":"2026-10-16T09:45:52Z","timestamp":1792143952502,"version":"4.4.0"},"reference-count":53,"publisher":"Wiley","issue":"5","license":[{"start":{"date-parts":[[2026,10,16]],"date-time":"2026-10-16T00:00:00Z","timestamp":1792108800000},"content-version":"vor","delay-in-days":45,"URL":"http:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0\/"},{"start":{"date-parts":[[2026,9,1]],"date-time":"2026-09-01T00:00:00Z","timestamp":1788220800000},"content-version":"tdm","delay-in-days":0,"URL":"http:\/\/doi.wiley.com\/10.1002\/tdm_license_1.1"}],"funder":[{"DOI":"10.13039\/100008398","name":"Villum Fonden","doi-asserted-by":"publisher","award":["00028156"],"award-info":[{"award-number":["00028156"]}],"id":[{"id":"10.13039\/100008398","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100008398","name":"Villum Fonden","doi-asserted-by":"publisher","award":["VIL60791"],"award-info":[{"award-number":["VIL60791"]}],"id":[{"id":"10.13039\/100008398","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100008398","name":"Villum Fonden","doi-asserted-by":"publisher","award":["VIL50371"],"award-info":[{"award-number":["VIL50371"]}],"id":[{"id":"10.13039\/100008398","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100004836","name":"Danmarks Frie Forskningsfond","doi-asserted-by":"publisher","award":["DFF\u20108022\u201000301B"],"award-info":[{"award-number":["DFF\u20108022\u201000301B"]}],"id":[{"id":"10.13039\/501100004836","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000936","name":"Gordon and Betty Moore Foundation","doi-asserted-by":"publisher","award":["GBMF9206"],"award-info":[{"award-number":["GBMF9206"]}],"id":[{"id":"10.13039\/100000936","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["onlinelibrary.wiley.com"],"crossmark-restriction":true},"short-container-title":["Physiologia Plantarum"],"published-print":{"date-parts":[[2026,9]]},"abstract":"ABSTRACT<\/jats:title>\n \n Seagrass photosynthesis is often considered carbon\u2010limited due to slow diffusion rates across leaf diffusive boundary layers (DBLs) and low CO\n 2<\/jats:sub>\n availability in seawater. Yet, seagrasses are important carbon sinks in coastal environments, enabled via CO\n 2<\/jats:sub>\n concentrating mechanisms (CCMs), allowing for bicarbonate (HCO\n 3<\/jats:sub>\n \u2212<\/jats:sup>\n ) utilization in their photosynthesis. However, the role of DBL acidification in seagrass CCMs is not well described. Here, we used pH\u2010sensitive planar optodes with micro\u2010crystalline diamond powder, pH and O\n 2<\/jats:sub>\n \u2010sensitive optical sensor nanoparticles, and microsensor measurements to determine the pH, O\n 2<\/jats:sub>\n , and inorganic carbon dynamics on the adaxial and abaxial domains of seagrass (\n \n Zostera marina<\/jats:italic>\n <\/jats:styled-content>\n L.) leaves. We provide first experimental evidence of seagrass leaf pH polarization via relative adaxial surface acidification, leading to enhanced CO\n 2<\/jats:sub>\n formation at the leaf surface via increased dehydration of HCO\n 3<\/jats:sub>\n \u2212<\/jats:sup>\n and carbonate equilibrium speciation shifts towards CO\n 2<\/jats:sub>\n . This enabled enhanced photosynthetic O\n 2<\/jats:sub>\n evolution and CO\n 2<\/jats:sub>\n flux to the adaxial domain of leaves (i.e., ~2\u2010fold higher CO\n 2<\/jats:sub>\n availability and flux as compared with the abaxial surface) under low light conditions and at the onset of photosynthesis during dark\/light transitions under saturating light. Such a CCM\u2010like mechanism for improved CO\n 2<\/jats:sub>\n formation around seagrass leaves can be vital in dense seagrass meadows with high diel fluctuations in water\u2010column light, pH, and O\n 2<\/jats:sub>\n concentrations to sustain adequate carbon assimilation.\n <\/jats:p>","DOI":"10.1111\/ppl.71158","type":"journal-article","created":{"date-parts":[[2026,10,16]],"date-time":"2026-10-16T09:03:44Z","timestamp":1792141424000},"update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Leaf\n pH<\/scp>\n Polarization in the Seagrass\n Zostera marina<\/i>\n Enhances Adaxial Surface\n \n CO\n 2<\/sub>\n <\/scp>\n Formation and Photosynthesis\u2010Driven Carbon Fixation"],"prefix":"10.1111","volume":"178","author":[{"ORCID":"https:\/\/orcid.org\/0000-0001-9010-1179","authenticated-orcid":false,"given":"Kasper Elgetti","family":"Brodersen","sequence":"first","affiliation":[{"name":"Marine Biological Section, Department of Biology University of Copenhagen Helsing\u00f8r Denmark"},{"name":"Environmental Dynamics Section, Department of Science and Environment Roskilde University Roskilde Denmark"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Maria","family":"Mosshammer","sequence":"additional","affiliation":[{"name":"Marine Biological Section, Department of Biology University of Copenhagen Helsing\u00f8r Denmark"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-9270-8863","authenticated-orcid":false,"given":"Zs\u00f3fia","family":"Bogn\u00e1r","sequence":"additional","affiliation":[{"name":"Department of Inorganic and Analytical Chemistry Budapest University of Technology and Economics Budapest Hungary"},{"name":"Center for Visualizing Catalytic Processes (VISION), Department of Physics Technical University of Denmark (DTU) Kongens Lyngby Denmark"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-1792-4790","authenticated-orcid":false,"given":"Michael","family":"K\u00fchl","sequence":"additional","affiliation":[{"name":"Marine Biological Section, Department of Biology University of Copenhagen Helsing\u00f8r Denmark"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"311","published-online":{"date-parts":[[2026,10,16]]},"reference":[{"issue":"10","key":"e_1_2_9_2_1","doi-asserted-by":"crossref","first-page":"2625","DOI":"10.1002\/ece3.333","article-title":"Effects of CO2 Enrichment on Photosynthesis, Growth, and Nitrogen Metabolism of the Seagrass \n Zostera noltii","volume":"2","author":"Alexandre A.","year":"2012","journal-title":"Ecology and Evolution"},{"key":"e_1_2_9_3_1","doi-asserted-by":"crossref","first-page":"59","DOI":"10.3354\/meps12439","article-title":"Response to Experimental Warming in Northern Eelgrass Populations: Comparison Across a Range of Temperature Adaptations","volume":"589","author":"Beca\u2010Carretero P.","year":"2018","journal-title":"Marine Ecology Progress Series"},{"issue":"3","key":"e_1_2_9_4_1","first-page":"349","article-title":"Inorganic Carbon Utilization in Marine Angiosperms (Seagrasses)","volume":"29","author":"Beer S.","year":"2002","journal-title":"Functional Plant Biology"},{"issue":"1","key":"e_1_2_9_5_1","doi-asserted-by":"crossref","first-page":"99","DOI":"10.1016\/0022-0981(83)90152-1","article-title":"Photosynthesis of Ulva sp. II. Utilization of CO2 and HCO3\n \u2212 When Submerged","volume":"70","author":"Beer S.","year":"1983","journal-title":"Journal of Experimental Marine Biology and Ecology"},{"issue":"3","key":"e_1_2_9_6_1","doi-asserted-by":"crossref","first-page":"277","DOI":"10.1016\/S0304-3770(96)01109-6","article-title":"The Acquisition of Inorganic Carbon by the Seagrass \n Zostera marina","volume":"56","author":"Beer S.","year":"1997","journal-title":"Aquatic Botany"},{"key":"e_1_2_9_7_1","doi-asserted-by":"crossref","first-page":"1763","DOI":"10.1021\/acssensors.3c01941","article-title":"Multiparameter Sensing of Oxygen and pH at Biological Interfaces via Hyperspectral Imaging of Luminescent Sensor Nanoparticles","volume":"9","author":"Bogn\u00e1r Z.","year":"2024","journal-title":"ACS Sensors"},{"key":"e_1_2_9_8_1","doi-asserted-by":"crossref","first-page":"52","DOI":"10.1016\/j.snb.2008.08.028","article-title":"Fluorescent Poly(Styrene\u2010Block\u2010Vinylpyrrolidone) Nanobeads for Optical Sensing of pH","volume":"139","author":"Borisov S. M.","year":"2009","journal-title":"Sensors and Actuators B: Chemical"},{"key":"e_1_2_9_9_1","doi-asserted-by":"crossref","first-page":"148","DOI":"10.1111\/j.1365-2745.2004.00943.x","article-title":"The Potential Role of Plant Oxygen and Sulphide Dynamics in Die\u2010Off Events of the Tropical Seagrass, \n Thalassia testudinum","volume":"93","author":"Borum J.","year":"2005","journal-title":"Journal of Ecology"},{"issue":"6","key":"e_1_2_9_10_1","doi-asserted-by":"crossref","first-page":"1240","DOI":"10.1111\/pce.12658","article-title":"Photosynthetic Response to Globally Increasing CO2 of CO\u2010Occurring Temperate Seagrass Species","volume":"39","author":"Borum J.","year":"2016","journal-title":"Plant, Cell & Environment"},{"key":"e_1_2_9_11_1","doi-asserted-by":"crossref","DOI":"10.3389\/fpls.2017.00657","article-title":"Sediment Resuspension and Deposition on Seagrass Leaves Impedes Internal Plant Aeration and Promotes Phytotoxic H2S Intrusion","volume":"8","author":"Brodersen K. E.","year":"2017","journal-title":"Frontiers in Plant Science"},{"key":"e_1_2_9_12_1","doi-asserted-by":"crossref","first-page":"1619","DOI":"10.1111\/pce.12740","article-title":"Nanoparticle\u2010Based Measurements of pH and O2 Dynamics in the Rhizosphere of \n Zostera marina\n L.: Effects of Temperature Elevation and Light\/Dark Transitions","volume":"39","author":"Brodersen K. E.","year":"2016","journal-title":"Plant, Cell & Environment"},{"issue":"24","key":"e_1_2_9_13_1","doi-asserted-by":"crossref","first-page":"14155","DOI":"10.1021\/acs.est.7b03878","article-title":"Seagrass\u2010Mediated Phosphorus and Iron Solubilization in Tropical Sediments","volume":"51","author":"Brodersen K. 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A.","year":"1999","journal-title":"Journal of Experimental Botany"},{"issue":"1","key":"e_1_2_9_23_1","doi-asserted-by":"crossref","first-page":"210","DOI":"10.4319\/lo.2003.48.1.0210","article-title":"Meristematic Oxygen Variability in Eelgrass (\n Zostera marina\n )","volume":"48","author":"Greve T. M.","year":"2003","journal-title":"Limnology and Oceanography"},{"key":"e_1_2_9_24_1","doi-asserted-by":"crossref","DOI":"10.3389\/fmars.2022.822485","article-title":"Temperature Effects on Leaf and Epiphyte Photosynthesis, Bicarbonate Use and Diel O2 Budgets of the Seagrass \n Zostera marina\n L","volume":"9","author":"Hansen A. B.","year":"2022","journal-title":"Frontiers in Marine Science"},{"issue":"1","key":"e_1_2_9_25_1","doi-asserted-by":"crossref","first-page":"55","DOI":"10.1016\/S0304-3770(00)00132-7","article-title":"A Buffer Sensitive Inorganic Carbon Utilization System in \n Zostera marina","volume":"69","author":"Hellblom F.","year":"2001","journal-title":"Aquatic Botany"},{"issue":"3","key":"e_1_2_9_26_1","doi-asserted-by":"crossref","first-page":"453","DOI":"10.1046\/j.1529-8817.2000.99139.x","article-title":"Water Motion, Marine Macroalgal Physiology, and Production","volume":"36","author":"Hurd C. L.","year":"2000","journal-title":"Journal of Phycology"},{"issue":"2","key":"e_1_2_9_27_1","doi-asserted-by":"crossref","first-page":"147","DOI":"10.1007\/s11120-017-0452-1","article-title":"Low Oxygen Affects Photophysiology and the Level of Expression of Two\u2010Carbon Metabolism Genes in the Seagrass Zostera muelleri","volume":"136","author":"Kim M.","year":"2018","journal-title":"Photosynthesis Research"},{"issue":"4","key":"e_1_2_9_28_1","doi-asserted-by":"crossref","first-page":"767","DOI":"10.1007\/BF00347527","article-title":"Hydrodynamics, Diffusion\u2010Boundary Layers and Photosynthesis of the Seagrasses Thalassia testudinum and Cymodocea nodosa","volume":"118","author":"Koch E.","year":"1994","journal-title":"Marine Biology"},{"issue":"1","key":"e_1_2_9_29_1","doi-asserted-by":"crossref","first-page":"68","DOI":"10.1080\/17451000510019015","article-title":"The Chemical Microenvironment of the Symbiotic Planktonic Foraminifer \n Orbulina universa","volume":"1","author":"K\u00f6hler\u2010Rink S.","year":"2005","journal-title":"Marine Biology Research"},{"issue":"4","key":"e_1_2_9_30_1","doi-asserted-by":"crossref","first-page":"2286","DOI":"10.1021\/es505734b","article-title":"Optical Sensor Nanoparticles in Artificial Sediments \u2013 A New Tool to Visualize O2 Dynamics Around the Rhizome and Roots of Seagrasses","volume":"49","author":"Koren K.","year":"2015","journal-title":"Environmental Science & Technology"},{"key":"e_1_2_9_31_1","doi-asserted-by":"crossref","first-page":"180","DOI":"10.1093\/oso\/9780195118810.003.0008","volume-title":"The Benthic Boundary Layer","author":"K\u00fchl M.","year":"2001"},{"issue":"14","key":"e_1_2_9_32_1","doi-asserted-by":"crossref","first-page":"3773","DOI":"10.1093\/jxb\/erx206","article-title":"Carbon\u2010Concentrating Mechanisms in Seagrasses","volume":"68","author":"Larkum A. W. D.","year":"2017","journal-title":"Journal of Experimental Botany"},{"key":"e_1_2_9_33_1","volume-title":"SEAGRASSES: BIOLOGY, ECOLOGYAND CONSERVATION","author":"Larkum A. W.","year":"2006"},{"key":"e_1_2_9_34_1","volume-title":"Biology of Seagrass","author":"Larkum A. W. D.","year":"1989"},{"key":"e_1_2_9_35_1","doi-asserted-by":"crossref","first-page":"315","DOI":"10.1007\/978-3-319-71354-0_11","volume-title":"Seagrasses of Australia","author":"Larkum A. W. D.","year":"2018"},{"key":"e_1_2_9_36_1","first-page":"348","article-title":"A Simple and Inexpensive High Resolution Color Ratiometric Planar Optode Imaging Approach: Application to Oxygen and pH Sensing","volume":"9","author":"Larsen M.","year":"2011","journal-title":"Limnology and Oceanography: Methods"},{"issue":"6","key":"e_1_2_9_37_1","doi-asserted-by":"crossref","first-page":"897","DOI":"10.4319\/lo.1973.18.6.0897","article-title":"Measurement of the Apparent Dissociation Constants of Carbonic Acid in Seawater at Atmospheric Pressure","volume":"18","author":"Mehrbach C.","year":"1973","journal-title":"Limnology and Oceanography"},{"issue":"2","key":"e_1_2_9_38_1","doi-asserted-by":"crossref","first-page":"126","DOI":"10.1007\/s00604-018-3202-y","article-title":"Nanoparticle\u2010 and Microparticle\u2010Based Luminescence Imaging of Chemical Species and Temperature in Aquatic Systems: A Review","volume":"186","author":"Mo\u00dfhammer M.","year":"2019","journal-title":"Microchimica Acta"},{"issue":"6","key":"e_1_2_9_39_1","doi-asserted-by":"crossref","first-page":"681","DOI":"10.1021\/acssensors.6b00071","article-title":"Design and Application of an Optical Sensor for Simultaneous Imaging of pH and Dissolved O2 With Low Cross\u2010Talk","volume":"1","author":"Mo\u00dfhammer M.","year":"2016","journal-title":"ACS Sensors"},{"key":"e_1_2_9_40_1","first-page":"1137","article-title":"Salt\u2010Tolerant ATPase Activity in the Plasmalemma of the Marine Angiosperm \n Zostera marina\n L","volume":"43","author":"Muramatsu Y.","year":"2002","journal-title":"Plant, Cell and Environment"},{"issue":"171","key":"e_1_2_9_41_1","doi-asserted-by":"crossref","DOI":"10.1098\/rsif.2020.0485","article-title":"Flow and Epiphyte Growth Effects on the Thermal, Optical and Chemical Microenvironment in the Leaf Phyllosphere of Seagrass (\n Zostera marina\n )","volume":"17","author":"Noisette F.","year":"2020","journal-title":"Journal of the Royal Society Interface"},{"key":"e_1_2_9_42_1","doi-asserted-by":"crossref","first-page":"1329","DOI":"10.1111\/1365-2435.13067","article-title":"Abiotic and Biotic Interactions in the Diffusive Boundary Layer of Kelp Blades Create a Potential Refuge From Ocean Acidification","volume":"32","author":"Noisette F.","year":"2018","journal-title":"Functional Ecology"},{"key":"e_1_2_9_43_1","doi-asserted-by":"crossref","DOI":"10.3389\/fpls.2013.00140","article-title":"Underwater Photosynthesis of Submerged Plants\u2013Recent Advances and Methods","volume":"4","author":"Pedersen O.","year":"2013","journal-title":"Frontiers in Plant Science"},{"issue":"5","key":"e_1_2_9_44_1","doi-asserted-by":"crossref","first-page":"818","DOI":"10.1104\/pp.66.5.818","article-title":"Photosynthetic HCO3\n \u2212 Utilization and OH\u2212 Excretion in Aquatic Angiosperms: Light\u2010Induced pH Changes at the Leaf Surface","volume":"66","author":"Prins H. B.","year":"1980","journal-title":"Plant Physiology"},{"key":"e_1_2_9_45_1","volume-title":"Energetics and Transport in Aquatic Plants","author":"Raven J. A.","year":"1984"},{"key":"e_1_2_9_46_1","first-page":"85","volume-title":"Advances in Botanical Research","author":"Raven J. A.","year":"1997"},{"issue":"2","key":"e_1_2_9_47_1","doi-asserted-by":"crossref","first-page":"111","DOI":"10.1007\/s11120-013-9962-7","article-title":"Energy Costs of Carbon Dioxide Concentrating Mechanisms in Aquatic Organisms","volume":"121","author":"Raven J. A.","year":"2014","journal-title":"Photosynthesis Research"},{"issue":"1","key":"e_1_2_9_48_1","doi-asserted-by":"crossref","first-page":"105","DOI":"10.1007\/s11120-012-9768-z","article-title":"Ecophysiology of Photosynthesis in Macroalgae","volume":"113","author":"Raven J. A.","year":"2012","journal-title":"Photosynthesis Research"},{"issue":"11","key":"e_1_2_9_49_1","doi-asserted-by":"crossref","first-page":"2580","DOI":"10.1111\/gcb.15594","article-title":"Coast\u2010Wide Evidence of Low pH Amelioration by Seagrass Ecosystems","volume":"27","author":"Ricart A. M.","year":"2021","journal-title":"Global Change Biology"},{"key":"e_1_2_9_50_1","doi-asserted-by":"crossref","first-page":"2820","DOI":"10.1111\/pce.13057","article-title":"Direct Uptake of HCO3\n \u2212 in the Marine Angiosperm \n Posidonia oceanica\n (L.) Delile Driven by a Plasma Membrane H+ Economy","volume":"40","author":"Rubio L.","year":"2017","journal-title":"Plant, Cell & Environment"},{"issue":"2","key":"e_1_2_9_51_1","doi-asserted-by":"crossref","first-page":"283","DOI":"10.1111\/j.1365-2427.1992.tb00540.x","article-title":"Photosynthetic Use of Inorganic Carbon Among Primary and Secondary Water Plants in Streams","volume":"27","author":"Sand\u2010Jensen K.","year":"1992","journal-title":"Freshwater Biology"},{"issue":"2","key":"e_1_2_9_52_1","doi-asserted-by":"crossref","first-page":"139","DOI":"10.1016\/j.jembe.2011.05.031","article-title":"Seasonal Acclimation in Metabolism Reduces Light Requirements of Eelgrass (\n Zostera marina\n )","volume":"407","author":"Staehr P. A.","year":"2011","journal-title":"Journal of Experimental Marine Biology and Ecology"},{"issue":"7","key":"e_1_2_9_53_1","doi-asserted-by":"crossref","first-page":"1180","DOI":"10.1093\/pcp\/pcs074","article-title":"Leaf Adaxial\u2013Abaxial Polarity Specification and Lamina Outgrowth: Evolution and Development","volume":"53","author":"Yamaguchi T.","year":"2012","journal-title":"Plant and Cell Physiology"},{"key":"e_1_2_9_54_1","article-title":"Effects of Epiphytic Biofilm Activity on the Photosynthetic Activity, pH and Inorganic Carbon Microenvironment of Seagrass Leaves (\n Zostera marina\n L.)","volume":"9","author":"Zhang Q.","year":"2022","journal-title":"Frontiers in Marine Science"}],"container-title":["Physiologia Plantarum"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/onlinelibrary.wiley.com\/doi\/pdf\/10.1111\/ppl.71158","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/onlinelibrary.wiley.com\/doi\/full-xml\/10.1111\/ppl.71158","content-type":"application\/xml","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/onlinelibrary.wiley.com\/doi\/pdf\/10.1111\/ppl.71158","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,10,16]],"date-time":"2026-10-16T09:03:51Z","timestamp":1792141431000},"score":1,"resource":{"primary":{"URL":"https:\/\/onlinelibrary.wiley.com\/doi\/10.1111\/ppl.71158"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,9]]},"references-count":53,"journal-issue":{"issue":"5","published-print":{"date-parts":[[2026,9]]}},"alternative-id":["10.1111\/ppl.71158"],"URL":"https:\/\/doi.org\/10.1111\/ppl.71158","archive":["Portico"],"relation":{},"ISSN":["0031-9317","1399-3054"],"issn-type":[{"value":"0031-9317","type":"print"},{"value":"1399-3054","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,9]]},"assertion":[{"value":"2026-08-13","order":0,"name":"received","label":"Received","group":{"name":"publication_history","label":"Publication History"}},{"value":"2026-10-07","order":2,"name":"accepted","label":"Accepted","group":{"name":"publication_history","label":"Publication History"}},{"value":"2026-10-16","order":3,"name":"published","label":"Published","group":{"name":"publication_history","label":"Publication History"}}],"article-number":"e71158"}}