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Although both are required for biomass accumulation and efficient photosynthesis, the chloroplast \u03b2CA1 does not directly participate in carbon assimilation but instead specifically coordinates photosystem performance and leaf internal structure. This role is distinct from the cytosol\/plasma\u2010membrane \u03b2CA2 that supports carbon assimilation and overall growth.\n                        <\/jats:p>\n                      <\/jats:list-item>\n                      <jats:list-item>\n                        <jats:p>\n                          We demonstrate that \u03b2CA1 buffers light\u2010induced stromal pH changes, thereby optimizing proton motive force (\n                          <jats:italic>pmf<\/jats:italic>\n                          ) and stabilizing photosystem II performance. The absence of \u03b2CA1 increases photosystem sensitivity to high\u2010light intensities, resulting in increased photoprotective responses and adaptive thylakoid re\u2010organization when environmental CO\n                          <jats:sub>2<\/jats:sub>\n                          becomes limiting relative to irradiance.\n                        <\/jats:p>\n                      <\/jats:list-item>\n                      <jats:list-item>\n                        <jats:p>\n                          Our findings establish chloroplast \u03b2CA1 as a key integrator of CO\n                          <jats:sub>2<\/jats:sub>\n                          and light cues that optimizes photosystem performance through modulation of\n                          <jats:italic>pmf<\/jats:italic>\n                          partitioning and reveal stromal pH regulation as a previously unrecognized mechanism for optimizing photosynthesis under suboptimal CO\n                          <jats:sub>2<\/jats:sub>\n                          and light conditions.\n                        <\/jats:p>\n                      <\/jats:list-item>\n                    <\/jats:list>\n                  <\/jats:p>","DOI":"10.1111\/nph.71578","type":"journal-article","created":{"date-parts":[[2026,9,18]],"date-time":"2026-09-18T03:40:23Z","timestamp":1789702823000},"update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["\u03b2\u2010Carbonic anhydrases integrate\n                    <scp>\n                      CO\n                      <sub>2<\/sub>\n                    <\/scp>\n                    and light cues to coordinate photosynthesis in tomato"],"prefix":"10.1111","author":[{"ORCID":"https:\/\/orcid.org\/0009-0000-8583-2575","authenticated-orcid":false,"given":"Anran","family":"Wang","sequence":"first","affiliation":[{"name":"Department of Horticulture, College of Agriculture and Biotechnology Zhejiang University  Hangzhou 310058 China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0008-3097-1648","authenticated-orcid":false,"given":"Chenxin","family":"Yue","sequence":"additional","affiliation":[{"name":"Department of Horticulture, College of Agriculture and Biotechnology Zhejiang University  Hangzhou 310058 China"},{"name":"Hainan Institute Zhejiang University, Yazhou Bay Science and Technology City  Sanya 572025 China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xiao","family":"Liang","sequence":"additional","affiliation":[{"name":"Department of Horticulture, College of Agriculture and Biotechnology Zhejiang University  Hangzhou 310058 China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0005-0662-744X","authenticated-orcid":false,"given":"Haiming","family":"Ying","sequence":"additional","affiliation":[{"name":"Department of Horticulture, College of Agriculture and Biotechnology Zhejiang University  Hangzhou 310058 China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jianrong","family":"Lv","sequence":"additional","affiliation":[{"name":"Department of Horticulture, College of Agriculture and Biotechnology Zhejiang University  Hangzhou 310058 China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Shujun","family":"Shao","sequence":"additional","affiliation":[{"name":"Department of Horticulture, College of Agriculture and Biotechnology Zhejiang University  Hangzhou 310058 China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Hongxin","family":"Wu","sequence":"additional","affiliation":[{"name":"Chinese Academy of Agricultural Mechanization Sciences Group Co. Ltd  Beijing 100083 China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Minghan","family":"Xu","sequence":"additional","affiliation":[{"name":"Chinese Academy of Agricultural Mechanization Sciences Group Co. Ltd  Beijing 100083 China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yang","family":"Liu","sequence":"additional","affiliation":[{"name":"Chinese Academy of Agricultural Mechanization Sciences Group Co. Ltd  Beijing 100083 China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xiaopeng","family":"Guo","sequence":"additional","affiliation":[{"name":"Chinese Academy of Agricultural Mechanization Sciences Group Co. Ltd  Beijing 100083 China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Wei","family":"Yuan","sequence":"additional","affiliation":[{"name":"Horgos Agricultural Science and Technology Development Group Co. Ltd  Yili 835200 China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-5989-6989","authenticated-orcid":false,"given":"Christine H.","family":"Foyer","sequence":"additional","affiliation":[{"name":"School of Biosciences, College of Life and Environmental, Sciences University of Birmingham  Edgbaston B15 2TT UK"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-5351-1910","authenticated-orcid":false,"given":"Kai","family":"Shi","sequence":"additional","affiliation":[{"name":"Department of Horticulture, College of Agriculture and Biotechnology Zhejiang University  Hangzhou 310058 China"},{"name":"Hainan Institute Zhejiang University, Yazhou Bay Science and Technology City  Sanya 572025 China"},{"name":"Key Laboratory of Horticultural Plants Growth and Development Ministry of Agriculture and Rural Affairs of China  Hangzhou 310058 China"},{"name":"Zhejiang Key Laboratory of Horticultural Crop Quality Improvement  Hangzhou 310058 China"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"311","published-online":{"date-parts":[[2026,9,17]]},"reference":[{"key":"e_1_2_9_2_1","doi-asserted-by":"crossref","first-page":"9123","DOI":"10.1073\/pnas.0800909105","article-title":"Intraplastidial trafficking of a phage\u2010type RNA polymerase is mediated by a thylakoid RING\u2010H2 protein","volume":"105","author":"Azevedo J","year":"2008","journal-title":"Proceedings of the National Academy of Sciences, USA"},{"key":"e_1_2_9_3_1","doi-asserted-by":"crossref","first-page":"2000","DOI":"10.1111\/nph.17839","article-title":"Chloroplast development in green plant tissues: the interplay between light, hormone, and transcriptional regulation","volume":"233","author":"Cackett L","year":"2022","journal-title":"New Phytologist"},{"key":"e_1_2_9_4_1","doi-asserted-by":"crossref","first-page":"1081","DOI":"10.1039\/b507310n","article-title":"Granal stacking of thylakoid membranes 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X","year":"2020","journal-title":"Plant Physiology"},{"key":"e_1_2_9_29_1","doi-asserted-by":"crossref","first-page":"841","DOI":"10.1007\/s00442-021-05062-y","article-title":"The limiting factors and regulatory processes that control the environmental responses of C3, C3\u2013C4 intermediate, and C4 photosynthesis","volume":"197","author":"Johnson JE","year":"2021","journal-title":"Oecologia"},{"key":"e_1_2_9_30_1","doi-asserted-by":"crossref","first-page":"1518","DOI":"10.1111\/pbi.13830","article-title":"Overexpression of cytoplasmic C4\n                  Flaveria bidentis carbonic anhydrase in C3\n                  Arabidopsis thaliana increases amino acids, photosynthetic potential, and biomass","volume":"20","author":"Kandoi D","year":"2022","journal-title":"Plant Biotechnology Journal"},{"key":"e_1_2_9_31_1","doi-asserted-by":"crossref","DOI":"10.1098\/rstb.2013.0225","article-title":"Structural changes of the thylakoid membrane network induced by high light stress in plant 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Procedures and sources of error","volume":"54","author":"Long SP","year":"2003","journal-title":"Journal of Experimental Botany"},{"key":"e_1_2_9_42_1","doi-asserted-by":"crossref","first-page":"56","DOI":"10.1016\/j.cell.2015.03.019","article-title":"Meeting the global food demand of the future by engineering crop photosynthesis and yield potential","volume":"161","author":"Long SP","year":"2015","journal-title":"Cell"},{"key":"e_1_2_9_43_1","doi-asserted-by":"crossref","first-page":"204","DOI":"10.1111\/nph.20429","article-title":"Alternative transcriptional initiation of OsbetaCA1 produces three distinct subcellular localization isoforms involved in stomatal response regulation and photosynthesis in rice","volume":"246","author":"Mao C","year":"2025","journal-title":"New Phytologist"},{"key":"e_1_2_9_44_1","doi-asserted-by":"crossref","first-page":"1688","DOI":"10.1093\/pcp\/pcn140","article-title":"Effect of PGR5 impairment on photosynthesis and growth in Arabidopsis 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