{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,21]],"date-time":"2026-09-21T12:15:40Z","timestamp":1789992940246,"version":"4.0.1"},"reference-count":121,"publisher":"Oxford University Press (OUP)","issue":"1","license":[{"start":{"date-parts":[[2026,4,29]],"date-time":"2026-04-29T00:00:00Z","timestamp":1777420800000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/academic.oup.com\/pages\/standard-publication-reuse-rights"}],"funder":[{"DOI":"10.13039\/100031064","name":"Hypothesis Fund","doi-asserted-by":"publisher","id":[{"id":"10.13039\/100031064","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000015","name":"U.S. Department of Energy","doi-asserted-by":"publisher","award":["DE-SC0024175"],"award-info":[{"award-number":["DE-SC0024175"]}],"id":[{"id":"10.13039\/100000015","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000001","name":"National Science Foundation","doi-asserted-by":"publisher","award":["MCB-2213840"],"award-info":[{"award-number":["MCB-2213840"]}],"id":[{"id":"10.13039\/100000001","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000001","name":"National Science Foundation","doi-asserted-by":"publisher","award":["MCB-2213841"],"award-info":[{"award-number":["MCB-2213841"]}],"id":[{"id":"10.13039\/100000001","id-type":"DOI","asserted-by":"publisher"}]},{"name":"Canada Research Chair","award":["950\u2013232698"],"award-info":[{"award-number":["950\u2013232698"]}],"id":[{"id":"https:\/\/ror.org\/0517h6h17","id-type":"ROR","asserted-by":"crossref"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2026,9,1]]},"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>Ribulose-1,5-bisphosphate carboxylase\/oxygenase (Rubisco) underpins nearly all primary production yet remains a slow, error-prone enzyme because it catalyzes both carboxylation and oxygenation, the latter initiating photorespiration and reducing net carbon gain. Many organisms mitigate these limitations not by improving Rubisco selectivity for CO2 directly, but by modifying its local environment using CO\u2082-concentrating mechanisms (CCMs). In algae, a prominent biophysical CCM strategy is the pyrenoid: a phase-separated, Rubisco-rich condensate coupled to bicarbonate transport, local carbonic anhydrase activity, and diffusion barriers that elevate CO\u2082 at Rubisco active sites. Although pyrenoids have been most intensively studied in algal models, a pyrenoid-based CCM has evolved independently in a single land-plant lineage\u2014the hornworts\u2014providing a powerful comparative system for understanding how chloroplast organization can be tuned to terrestrial CO\u2082-delivery constraints. Here we synthesize a century of hornwort pyrenoid research in ecological, phylogenetic, and mechanistic context. We summarize bryophyte anatomical and microhabitat features that impose strong CO\u2082 diffusion limitation, and compare hornwort and algal pyrenoids in ultrastructure, molecular parts lists, and regulation. We highlight emerging models for hornwort pyrenoid formation, inorganic-carbon delivery, CO\u2082 generation and recapture, and review recent biochemical and structural work that has revealed distinctive hornwort Rubisco properties and biogenesis. Finally, we discuss how hornwort pyrenoids complement efforts to engineer algal pyrenoid components into C\u2083 crops, and propose modular, hybrid engineering strategies that leverage hornwort compatibility with the embryophyte chloroplast, while selectively importing algal modules. Together, hornwort pyrenoids illustrate both the convergent logic and the lineage-specific solutions of biophysical CO\u2082 concentration, and they open avenues for mechanistic discovery and photosynthesis engineering.<\/jats:p>","DOI":"10.1093\/plphys\/kiag247","type":"journal-article","created":{"date-parts":[[2026,4,23]],"date-time":"2026-04-23T11:44:01Z","timestamp":1776944641000},"source":"Crossref","is-referenced-by-count":2,"title":["Hornwort pyrenoids: a terrestrial exception with engineering lessons"],"prefix":"10.1093","volume":"202","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-1629-9786","authenticated-orcid":false,"given":"Tanner A","family":"Robison","sequence":"first","affiliation":[{"name":"Plant Biology Section, School of Integrative Plant Science, Cornell University , Ithaca, NY ,","place":["United States"]},{"name":"Boyce Thompson Institute , Ithaca, NY ,","place":["United States"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0770-1446","authenticated-orcid":false,"given":"Juan Carlos","family":"Villarreal A","sequence":"additional","affiliation":[{"name":"D\u00e9partement de Biologie, Universit\u00e9 Laval \u00a0 Qu\u00e9bec City, QC G1V 0A6 ,","place":["Canada"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0076-0152","authenticated-orcid":false,"given":"Fay-Wei","family":"Li","sequence":"additional","affiliation":[{"name":"Plant Biology Section, School of Integrative Plant Science, Cornell University , Ithaca, NY ,","place":["United States"]},{"name":"Boyce Thompson Institute , Ithaca, NY ,","place":["United States"]},{"name":"Department of Biology, Duke University , Durham, NC 27708 ,","place":["United States"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-2072-0884","authenticated-orcid":false,"given":"Laura H","family":"Gunn","sequence":"additional","affiliation":[{"name":"Plant Biology Section, School of Integrative Plant Science, Cornell University , Ithaca, NY ,","place":["United States"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"286","published-online":{"date-parts":[[2026,4,29]]},"reference":[{"key":"2026092107332686700_kiag247-B1","doi-asserted-by":"publisher","author":"Adler","year":"2025  18.","DOI":"10.1101\/2025.11.18.689020"},{"key":"2026092107332686700_kiag247-B2","doi-asserted-by":"publisher","first-page":"1272","DOI":"10.1126\/science.aap9221","article-title":"Plant RuBisCo assembly in E. coli with five chloroplast chaperones including 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