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Natl. Acad. Sci. U.S.A."],"published-print":{"date-parts":[[2026,9,15]]},"abstract":"<jats:p>\n                    Karrikins (KARs) are a class of butenolide molecules discovered in smoke hypothesized to mimic an undiscovered plant hormone, KAI2 ligand (KL). KAR\/KL signaling regulates germination, seedling development, stress tolerance, and symbiotic interactions with soil microbes, among other traits. KAR\/KL signaling is initiated by KARRIKIN INSENSITIVE2 (KAI2), an \u0251\/\u03b2-hydrolase related to the strigolactone enzyme-receptor DWARF14 (D14). Activated KAI2 forms protein\u2013protein interactions that trigger proteasomal degradation of a transcriptional regulator, SUPPRESSOR OF MAX2 1 (SMAX1), initiating changes in gene expression.\n                    <jats:italic toggle=\"yes\">D14-LIKE2<\/jats:italic>\n                    (\n                    <jats:italic toggle=\"yes\">DLK2<\/jats:italic>\n                    ), an ancient paralog of\n                    <jats:italic toggle=\"yes\">KAI2<\/jats:italic>\n                    and\n                    <jats:italic toggle=\"yes\">D14<\/jats:italic>\n                    , is a prominent transcriptional marker of KAR\/KL signaling in many plants that has uncertain function. We find that DLK2 forms a negative feedback loop that attenuates KAR\/KL signaling in\n                    <jats:italic toggle=\"yes\">Arabidopsis thaliana<\/jats:italic>\n                    . This mechanism complements that of\n                    <jats:italic toggle=\"yes\">KARRIKIN UPREGULATED F-BOX1<\/jats:italic>\n                    (\n                    <jats:italic toggle=\"yes\">KUF1<\/jats:italic>\n                    ), which putatively restricts KAR\/KL metabolism through targeted protein degradation. Loss-of-function mutations of\n                    <jats:italic toggle=\"yes\">DLK2<\/jats:italic>\n                    show little effect alone, but synthetically enhance the constitutive KAR\/KL responses of\n                    <jats:italic toggle=\"yes\">kuf1<\/jats:italic>\n                    seedlings. Overexpression of DLK2 proteins from several plants increases the abundance of a SMAX1 ratiometric reporter. DLK2 does not require nuclear localization to protect SMAX1, suggesting its function is independent of interactions with SMAX1 or its transcriptional regulator partners. DLK2 hydrolyzes a profluorescent, desmethyl butenolide reporter molecule that is putatively analogous to KL. We hypothesize that DLK2 catabolizes KAI2 ligand(s) without participating in KAR\/KL signaling directly. This functional antagonism could have evolved after\n                    <jats:italic toggle=\"yes\">KAI2<\/jats:italic>\n                    gene duplication through subfunctionalizing mutations that disrupted protein\u2013protein interactions while preserving enzymatic activity.\n                  <\/jats:p>","DOI":"10.1073\/pnas.2525145123","type":"journal-article","created":{"date-parts":[[2026,9,8]],"date-time":"2026-09-08T17:47:37Z","timestamp":1788889657000},"update-policy":"https:\/\/doi.org\/10.1073\/pnas.cm10313","source":"Crossref","is-referenced-by-count":1,"title":["Negative feedback regulation of karrikin signaling in\n                    <i>Arabidopsis thaliana<\/i>\n                    by an antagonistic paralog of karrikin receptors"],"prefix":"10.1073","volume":"123","author":[{"ORCID":"https:\/\/orcid.org\/0000-0001-5217-9813","authenticated-orcid":false,"given":"Qingtian","family":"Li","sequence":"first","affiliation":[{"id":[{"id":"https:\/\/ror.org\/03nawhv43","id-type":"ROR","asserted-by":"publisher"}],"name":"Department of Botany and Plant Sciences, University of California","place":["Riverside"]},{"name":"Yazhouwan National Laboratory","place":["Sanya, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-6196-4098","authenticated-orcid":false,"given":"Sun Hyun","family":"Chang","sequence":"additional","affiliation":[{"id":[{"id":"https:\/\/ror.org\/03nawhv43","id-type":"ROR","asserted-by":"publisher"}],"name":"Department of Botany and Plant Sciences, University of 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C. Nelson Feedback regulation of karrikin signaling by KUF1 and DLK2 in Arabidopsis thaliana. NCBI BioProject. https:\/\/www.ncbi.nlm.nih.gov\/bioproject\/1320694. Deposited 11 October 2025."}],"container-title":["Proceedings of the National Academy of Sciences"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/pnas.org\/doi\/pdf\/10.1073\/pnas.2525145123","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,9,8]],"date-time":"2026-09-08T17:48:28Z","timestamp":1788889708000},"score":1,"resource":{"primary":{"URL":"https:\/\/pnas.org\/doi\/10.1073\/pnas.2525145123"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,9,8]]},"references-count":101,"journal-issue":{"issue":"37","published-print":{"date-parts":[[2026,9,15]]}},"alternative-id":["10.1073\/pnas.2525145123"],"URL":"https:\/\/doi.org\/10.1073\/pnas.2525145123","relation":{},"ISSN":["0027-8424","1091-6490"],"issn-type":[{"value":"0027-8424","type":"print"},{"value":"1091-6490","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,9,8]]},"assertion":[{"value":"2025-09-15","order":0,"name":"received","label":"Received","group":{"name":"publication_history","label":"Publication History"}},{"value":"2026-07-30","order":2,"name":"accepted","label":"Accepted","group":{"name":"publication_history","label":"Publication History"}},{"value":"2026-09-08","order":3,"name":"published","label":"Published","group":{"name":"publication_history","label":"Publication History"}}],"article-number":"e2525145123"}}