{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,8,20]],"date-time":"2026-08-20T06:23:34Z","timestamp":1787207014517,"version":"build-2736575974"},"reference-count":57,"publisher":"Frontiers Media SA","license":[{"start":{"date-parts":[[2026,8,20]],"date-time":"2026-08-20T00:00:00Z","timestamp":1787184000000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100004837","name":"Ministerio de Ciencia e Innovaci\u00f3n","doi-asserted-by":"publisher","id":[{"id":"10.13039\/501100004837","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["frontiersin.org"],"crossmark-restriction":true},"short-container-title":["Front. Plant Sci."],"abstract":"\n Introduction<\/jats:title>\n \n Carotenoid cleavage dioxygenase 4 (\n CCD4<\/jats:italic>\n ) enzymes play central roles in carotenoid turnover and apocarotenoid biosynthesis in plants. Despite their importance, the evolutionary mechanisms underlying diversification of\n CCD4<\/jats:italic>\n catalytic functions remain poorly understood. This study investigated the CCD gene family in\n Buddleja alternifolia<\/jats:italic>\n , with particular emphasis on the expansion and functional evolution of the\n CCD4<\/jats:italic>\n subfamily.\n <\/jats:p>\n <\/jats:sec>\n \n Methods<\/jats:title>\n \n A genome-wide identification and comparative analysis of CCD genes were performed in\n B. alternifolia<\/jats:italic>\n . Genomic organization, phylogenetic relationships, and syntenic patterns were analyzed to investigate gene family expansion. Functional characterization of 11 Ba\n CCD4<\/jats:italic>\n paralogs was conducted through biochemical assays, while structural analyses were used to identify sequence features associated with differences in substrate cleavage specificity. Gene expression profiling was performed to assess patterns of tissue-specific regulation.\n <\/jats:p>\n <\/jats:sec>\n \n Results<\/jats:title>\n \n Twenty-three CCD genes were identified, including 12\n CCD4<\/jats:italic>\n paralogs, representing one of the largest\n CCD4<\/jats:italic>\n expansions reported within Lamiales. Syntenic and genomic analyses revealed that recent tandem duplication events, particularly within a\n CCD4<\/jats:italic>\n -rich region on chromosome 10, were the primary drivers of this expansion. The presence of pseudogenes in the same region supported an ongoing birth-and-death evolutionary process. Functional analyses demonstrated extensive biochemical diversification among Ba\n CCD4<\/jats:italic>\n enzymes despite their high sequence similarity. Several paralogs catalyzed asymmetric carotenoid cleavage leading to citraurin production, whereas two paralogs, KAG8367281 and KAG8375220, exhibited symmetric zeaxanthin cleavage activity, producing crocetin dialdehyde, the direct precursor of crocins. Notably, these crocetin-producing enzymes belonged to closely related paralogous pairs whose counterparts displayed distinct cleavage specificities, indicating rapid neofunctionalization after duplication. Structural analyses suggested that subtle sequence variations, including indels affecting loop regions adjacent to the substrate access channel, may underlie changes in regioselectivity. Expression profiling further revealed tissue-specific expression patterns consistent with functional divergence among paralogs.\n <\/jats:p>\n <\/jats:sec>\n \n Discussion<\/jats:title>\n \n These findings indicate that crocetin-forming activity in\n B. alternifolia<\/jats:italic>\n likely evolved through progressive modifications of ancestral\n CCD4<\/jats:italic>\n functions rather than through a single evolutionary event. The remarkable expansion and diversification of the\n CCD4<\/jats:italic>\n subfamily provide evidence for the role of gene duplication and neofunctionalization in shaping carotenoid cleavage specificity. Collectively, this work establishes\n B. alternifolia<\/jats:italic>\n as a valuable model for investigating the molecular evolution of\n CCD4<\/jats:italic>\n enzymes and the emergence of specialized apocarotenoid metabolism in plants.\n <\/jats:p>\n <\/jats:sec>","DOI":"10.3389\/fpls.2026.1916011","type":"journal-article","created":{"date-parts":[[2026,8,20]],"date-time":"2026-08-20T05:26:09Z","timestamp":1787203569000},"update-policy":"https:\/\/doi.org\/10.3389\/crossmark-policy","source":"Crossref","is-referenced-by-count":0,"title":["Evolutionary adaptation of CCD4 enzymes in Buddleja alternifolia for crocetin biosynthesis"],"prefix":"10.3389","volume":"17","author":[{"given":"Eduardo","family":"Parre\u00f1o","sequence":"first","affiliation":[{"name":"Instituto Universitario de Biotecnolog\u00eda, Agronom\u00eda y Sostenibilidad de la Universidad de Castilla-La Mancha (IBAS)","place":["Albacete, Spain"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Luc\u00eda","family":"Morote","sequence":"additional","affiliation":[{"name":"Instituto Universitario de Biotecnolog\u00eda, Agronom\u00eda y Sostenibilidad de la Universidad de Castilla-La Mancha (IBAS)","place":["Albacete, Spain"]},{"name":"Universit\u00e9 Jean Monnet Saint-Etienne, Centre National de la Recherche Scientifique, Laboratoire de Biotechnologies V\u00e9g\u00e9tales Appliqu\u00e9es aux Plantes Aromatiques et M\u00e9dicinales, Unit\u00e9 Mixte de Recherche","place":["Saint-Etienne, France"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Alberto J.","family":"L\u00f3pez Jim\u00e9nez","sequence":"additional","affiliation":[{"name":"Instituto Universitario de Biotecnolog\u00eda, Agronom\u00eda y Sostenibilidad de la Universidad de Castilla-La Mancha (IBAS)","place":["Albacete, Spain"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Elena","family":"Moreno-Gim\u00e9nez","sequence":"additional","affiliation":[{"name":"Instituto Universitario de Biotecnolog\u00eda, Agronom\u00eda y Sostenibilidad de la Universidad de Castilla-La Mancha (IBAS)","place":["Albacete, Spain"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"\u00c1ngela","family":"Rubio-Moraga","sequence":"additional","affiliation":[{"name":"Instituto Universitario de Biotecnolog\u00eda, Agronom\u00eda y Sostenibilidad de la Universidad de Castilla-La Mancha (IBAS)","place":["Albacete, Spain"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Oussama","family":"Ahrazem","sequence":"additional","affiliation":[{"name":"Instituto Universitario de Biotecnolog\u00eda, Agronom\u00eda y Sostenibilidad de la Universidad de Castilla-La Mancha (IBAS)","place":["Albacete, Spain"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Lourdes","family":"G\u00f3mez-G\u00f3mez","sequence":"additional","affiliation":[{"name":"Instituto Universitario de Biotecnolog\u00eda, Agronom\u00eda y Sostenibilidad de la Universidad de Castilla-La Mancha (IBAS)","place":["Albacete, Spain"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1965","published-online":{"date-parts":[[2026,8,20]]},"reference":[{"key":"B1","doi-asserted-by":"publisher","first-page":"14","DOI":"10.1093\/jxb\/erx277","article-title":"Evolutionarily distinct carotenoid cleavage dioxygenases are responsible for crocetin production in Buddleja davidii","volume":"68","author":"Ahrazem","year":"2017","journal-title":"J. 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