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In more than 60 C\n                    <jats:sub>4<\/jats:sub>\n                    lineages, CO\n                    <jats:sub>2<\/jats:sub>\n                    fixation is compartmentalized between tissues, and bundle-sheath cells become photosynthetically activated\n                    <jats:sup>2<\/jats:sup>\n                    . How the bundle sheath acquires this alternate identity that allows efficient photosynthesis is unclear. Here we show that changes to bundle-sheath gene expression in C\n                    <jats:sub>4<\/jats:sub>\n                    leaves are associated with the gain of a pre-existing\n                    <jats:italic>cis<\/jats:italic>\n                    -code found in the C\n                    <jats:sub>3<\/jats:sub>\n                    leaf. From single-nucleus gene-expression and chromatin-accessibility atlases, we uncover DNA binding with one finger (DOF) motifs that define bundle-sheath identity in the major crops C\n                    <jats:sub>3<\/jats:sub>\n                    rice and C\n                    <jats:sub>4<\/jats:sub>\n                    sorghum. Photosynthesis genes that are rewired to be strongly expressed in the bundle-sheath cells of C\n                    <jats:sub>4<\/jats:sub>\n                    sorghum acquire\n                    <jats:italic>cis<\/jats:italic>\n                    -elements that are recognized by DOFs. Our findings are consistent with a simple model in which C\n                    <jats:sub>4<\/jats:sub>\n                    photosynthesis is based on the recruitment of an ancestral\n                    <jats:italic>cis<\/jats:italic>\n                    -code associated with bundle-sheath identity. Gain of such elements harnessed a stable patterning of transcription factors between cell types that are found in both C\n                    <jats:sub>3<\/jats:sub>\n                    and C\n                    <jats:sub>4<\/jats:sub>\n                    leaves to activate photosynthesis in the bundle sheath. Our findings provide molecular insights into the evolution of the complex C\n                    <jats:sub>4<\/jats:sub>\n                    pathway, and might also guide the rational engineering of C\n                    <jats:sub>4<\/jats:sub>\n                    photosynthesis in C\n                    <jats:sub>3<\/jats:sub>\n                    crops to improve crop productivity and resilience\n                    <jats:sup>3,4<\/jats:sup>\n                    .\n                  <\/jats:p>","DOI":"10.1038\/s41586-024-08204-3","type":"journal-article","created":{"date-parts":[[2024,11,20]],"date-time":"2024-11-20T11:04:21Z","timestamp":1732100661000},"page":"143-150","update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":94,"title":["Exaptation of ancestral cell-identity networks enables C4 photosynthesis"],"prefix":"10.1038","volume":"636","author":[{"given":"Joseph","family":"Swift","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-7353-2556","authenticated-orcid":false,"given":"Leonie H.","family":"Luginbuehl","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Lei","family":"Hua","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-4440-1776","authenticated-orcid":false,"given":"Tina B.","family":"Schreier","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0006-9907-7762","authenticated-orcid":false,"given":"Ruth M.","family":"Donald","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Susan","family":"Stanley","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Na","family":"Wang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-8352-8956","authenticated-orcid":false,"given":"Travis A.","family":"Lee","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-0153-5659","authenticated-orcid":false,"given":"Joseph R.","family":"Nery","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-5799-5895","authenticated-orcid":false,"given":"Joseph R.","family":"Ecker","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-0662-7958","authenticated-orcid":false,"given":"Julian M.","family":"Hibberd","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2024,11,20]]},"reference":[{"key":"8204_CR1","doi-asserted-by":"crossref","unstructured":"Ghannoum, O., Evans, J. 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