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While the upper ocean was oxygenated, the deeper ocean was euxinic or ferruginous. Primary production was performed by Chlorobia, Cyanobacteria, Proteobacteria, and Archaeplastida. Cyanobacteria and Archaeplastida presumably performed oxygenic photosynthesis above the chemocline, while photosynthesis below the chemocline involved anoxygenic photolithotrophic oxidation of S\n                    <jats:sup>2\u2212<\/jats:sup>\n                    or Fe\n                    <jats:sup>2+<\/jats:sup>\n                    by Proteobacteria and S\n                    <jats:sup>2\u2212<\/jats:sup>\n                    by Chlorobia and Cyanobacteria. Photolithotrophic growth of extant Proteobacteria with Rubisco, and Chlorobia with the reverse tricarboxylic acid cycle, involves diffusive CO\n                    <jats:sub>2<\/jats:sub>\n                    entry; this likely also occurred under high Mesoproterozoic CO\n                    <jats:sub>2<\/jats:sub>\n                    . By contrast, Cyanobacteria may already have possessed CO\n                    <jats:sub>2<\/jats:sub>\n                    \u2010concentrating mechanisms comparable to extant lineages. Here, we evaluate how differences in primary carboxylation pathways and inorganic carbon acquisition among these groups influence stable carbon isotope fractionation and assess how isotopic signatures in modern representatives inform interpretations of Mesoproterozoic carbon isotope records. Although culture\u2010based fractionation patterns broadly overlap with Mesoproterozoic sedimentary \u03b4\n                    <jats:sup>13<\/jats:sup>\n                    C values, these records are strongly biased towards continental margin settings, and direct evidence for open\u2010ocean planktonic primary producers is lacking. The evidence reviewed highlights that physiological constraints on carbon acquisition and metabolism provide a framework for interpreting Mesoproterozoic carbon isotope signals with implications for primary production and the biological pump.\n                  <\/jats:p>","DOI":"10.1111\/nph.71181","type":"journal-article","created":{"date-parts":[[2026,4,30]],"date-time":"2026-04-30T05:36:29Z","timestamp":1777527389000},"page":"64-80","update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Photosynthetic primary production in the Mesoproterozoic"],"prefix":"10.1111","volume":"251","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-5216-2664","authenticated-orcid":false,"given":"Patricia","family":"S\u00e1nchez\u2010Baracaldo","sequence":"first","affiliation":[{"name":"School of Geographical Sciences University of Bristol  Bristol BS8 1SS UK"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-2789-3297","authenticated-orcid":false,"given":"John A.","family":"Raven","sequence":"additional","affiliation":[{"name":"Division of Plant Science University of Dundee at the James Hutton Institute  Invergowrie Dundee DD2 5DA UK"},{"name":"University of Technology, Sydney, Climate Change Cluster Faculty of Science  Sydney, Ultimo NSW 2007 Australia"},{"name":"School of Biological Sciences University of Western Australia  Crawley WA 6009 Australia"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"311","published-online":{"date-parts":[[2026,4,29]]},"reference":[{"key":"e_1_2_12_2_1","doi-asserted-by":"crossref","DOI":"10.1128\/aem.00863-24","article-title":"Evidence for autotrophic growth of purple sulfur bacteria using pyrite as electron and sulfur source","volume":"90","author":"Alarcon HV","year":"2024","journal-title":"Applied and Environmental Microbiology"},{"key":"e_1_2_12_3_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.bpj.2020.09.023"},{"key":"e_1_2_12_4_1","doi-asserted-by":"publisher","DOI":"10.1139\/b98-074"},{"key":"e_1_2_12_5_1","doi-asserted-by":"publisher","DOI":"10.1093\/jxb\/erm297"},{"key":"e_1_2_12_6_1","doi-asserted-by":"publisher","DOI":"10.1093\/jxb\/eri286"},{"key":"e_1_2_12_7_1","doi-asserted-by":"publisher","DOI":"10.1093\/jxb\/err417"},{"key":"e_1_2_12_8_1","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.1816654116"},{"key":"e_1_2_12_9_1","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.0503674102"},{"key":"e_1_2_12_10_1","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pbio.2000735"},{"key":"e_1_2_12_11_1","doi-asserted-by":"publisher","DOI":"10.1093\/sysbio\/syae025"},{"key":"e_1_2_12_12_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1472-4669.2009.00220.x"},{"key":"e_1_2_12_13_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-021-24396-y"},{"key":"e_1_2_12_14_1","doi-asserted-by":"publisher","DOI":"10.1038\/s42003-025-09247-6"},{"key":"e_1_2_12_15_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.tplants.2022.09.009"},{"key":"e_1_2_12_16_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.freeradbiomed.2019.05.004"},{"key":"e_1_2_12_17_1","doi-asserted-by":"publisher","DOI":"10.1038\/nature23457"},{"key":"e_1_2_12_18_1","doi-asserted-by":"publisher","DOI":"10.1038\/nature04068"},{"key":"e_1_2_12_19_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.gca.2007.12.006"},{"key":"e_1_2_12_20_1","doi-asserted-by":"publisher","DOI":"10.1134\/S002626171006010X"},{"key":"e_1_2_12_21_1","doi-asserted-by":"publisher","DOI":"10.1111\/1462-2920.14328"},{"key":"e_1_2_12_22_1","article-title":"Draft genome sequence of Chlorobium sp. 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