{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,23]],"date-time":"2026-09-23T19:02:19Z","timestamp":1790190139144,"version":"4.1.0"},"reference-count":49,"publisher":"Oxford University Press (OUP)","issue":"1","license":[{"start":{"date-parts":[[2025,10,27]],"date-time":"2025-10-27T00:00:00Z","timestamp":1761523200000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/academic.oup.com\/pages\/standard-publication-reuse-rights"}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2026,1,19]]},"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>The orchid perianth typically consists of three sepals and three petals, with the median petal transformed into a distinct structure lip. In our previously found P code model in orchids, A\/E protein OAGL6 interacts with B proteins to form L (OAGL6-2\/OAP3-2\/OPI) and SP (OAGL6-1\/OAP3-1\/OPI) complexes, which determine and sepal\/petal identities. This study demonstrates that OAGL6-1 and OAGL6-2 function redundantly with the E-class genes PaSEP1\/3 in regulating sepal\/petal and lip identities. Single-virus-induced gene silencing (VIGS) of OAGL6-1 resulted in mild morphological alterations in sepals\/petals, while silencing PaSEP1 or PaSEP3 individually caused little to no effect. However, triple silencing of OAGL6-1\/PaSEP1\/PaSEP3 led to additive and severe phenotypes, including greenish sepals\/petals with leaf-like epidermal features, indicating a collapse of the SP complex. Similarly, triple silencing of OAGL6-2\/PaSEP1\/PaSEP3 resulted in the transformation of the lip into a leaf-like structure, demonstrating their redundant roles in lip specification through the advanced L complex. Protein interaction assays confirmed strong interactions between OAGL6-1\/2 and PaSEP1\/3, supporting the formation of novel, advanced SP and L complexes. Additionally, PaSEP1 and PaSEP3 contribute to pedicel abscission and anthocyanin pigmentation, although their effects are weaker compared to OAGL6-1. Collectively, these findings highlight OAGL6-1 and OAGL6-2 as central regulators of perianth identity through interactions with PaSEP1\/3 in the SP and L complexes, respectively. PaSEP1\/3 function as minor yet essential partners that help maintain distinct floral organ identities and prevent reversion to leaf-like characteristics. This work expands the P code model and deepens our understanding of the unique mechanisms governing perianth specification in orchids.<\/jats:p>","DOI":"10.1093\/pcp\/pcaf137","type":"journal-article","created":{"date-parts":[[2025,10,23]],"date-time":"2025-10-23T12:25:08Z","timestamp":1761222308000},"page":"67-81","source":"Crossref","is-referenced-by-count":2,"title":["OAGL6 and PaSEP1\/3 cooperate in SP and L complexes of P code model to specify perianth identity in\n                    <i>Phalaenopsis<\/i>\n                    orchids"],"prefix":"10.1093","volume":"67","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-2824-9639","authenticated-orcid":false,"given":"Hsing-Fun","family":"Hsu","sequence":"first","affiliation":[{"name":"Institute of Biotechnology, National Chung Hsing University , Taichung , \u00a0 40227 ROC","place":["Taiwan"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Tzu-Chuan","family":"Hsiao","sequence":"additional","affiliation":[{"name":"Institute of Biotechnology, National Chung Hsing University , Taichung , \u00a0 40227 ROC","place":["Taiwan"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Wei-Han","family":"Hsu","sequence":"additional","affiliation":[{"name":"Institute of Biotechnology, National Chung Hsing University , Taichung , \u00a0 40227 ROC","place":["Taiwan"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Chen-Mao","family":"Liu","sequence":"additional","affiliation":[{"name":"Institute of Biotechnology, National Chung Hsing University , Taichung , \u00a0 40227 ROC","place":["Taiwan"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yi-Hsuan","family":"Shen","sequence":"additional","affiliation":[{"name":"Institute of Biotechnology, National Chung Hsing University , Taichung , \u00a0 40227 ROC","place":["Taiwan"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-2793-0262","authenticated-orcid":false,"given":"Chang-Hsien","family":"Yang","sequence":"additional","affiliation":[{"name":"Institute of Biotechnology, National Chung Hsing University , Taichung , \u00a0 40227 ROC","place":["Taiwan"]},{"name":"Advanced Plant and Food Crop Biotechnology Center, National Chung Hsing University , Taichung , \u00a0 40227 ROC","place":["Taiwan"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"286","published-online":{"date-parts":[[2025,10,27]]},"reference":[{"key":"2026021605165387800_ref1","first-page":"1","article-title":"Copper enzymes in isolated chloroplasts","volume":"24","author":"Arnon","year":"1949","journal-title":"Polyphenoloxidase in Beta vulgaris. Plant Physiol."},{"key":"2026021605165387800_ref2","doi-asserted-by":"crossref","first-page":"837","DOI":"10.1104\/pp.109.147116","article-title":"Characterization of the possible roles for B class MADS box genes in regulation of perianth formation in orchid","volume":"152","author":"Chang","year":"2010","journal-title":"Plant Physiol."},{"key":"2026021605165387800_ref3","doi-asserted-by":"crossref","first-page":"941","DOI":"10.1093\/jxb\/err323","article-title":"Functional analysis reveals the possible role of the C-terminal sequences and PI motif in the function of lily (Lilium longiflorum) PISTILLATA (PI) ortholog","volume":"63","author":"Chen","year":"2012","journal-title":"J. Exp. Bot."},{"key":"2026021605165387800_ref4","doi-asserted-by":"crossref","first-page":"31","DOI":"10.1038\/353031a0","article-title":"The war of the whorls: genetic 25 interactions controlling flower development","volume":"353","author":"Coen","year":"1991","journal-title":"Nature"},{"key":"2026021605165387800_ref5","doi-asserted-by":"crossref","first-page":"1792","DOI":"10.1093\/nar\/gkh340","article-title":"MUSCLE: multiple sequence alignment with high accuracy and high throughput","volume":"32","author":"Edgar","year":"2004","journal-title":"Nucleic Acids Res."},{"key":"2026021605165387800_ref6","doi-asserted-by":"crossref","first-page":"1107","DOI":"10.1093\/jexbot\/51.347.1107","article-title":"Functional role of anthocyanins in the leaves of Quintinia serrata A","volume":"51","author":"Gould","year":"2000","journal-title":"Cunn. J. Exp. Bot."},{"key":"2026021605165387800_ref7","doi-asserted-by":"crossref","first-page":"110","DOI":"10.1270\/jsbbs.60.110","article-title":"The expression of two DEFICIENS-like genes was reduced in the sepaloid tepals of viridiflora tulips","volume":"60","author":"Hirai","year":"2010","journal-title":"Breeding Sci."},{"key":"2026021605165387800_ref8","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1111\/tpj.12363","article-title":"AGAMOUS-LIKE13, a putative ancestor for the E functional genes, specifies male and female gametophyte morphogenesis","volume":"77","author":"Hsu","year":"2014","journal-title":"Plant J."},{"key":"2026021605165387800_ref9","doi-asserted-by":"crossref","first-page":"15046","DOI":"10.1038\/nplants.2015.46","article-title":"Model for perianth formation in orchids","volume":"1","author":"Hsu","year":"2015","journal-title":"Nat Plants"},{"key":"2026021605165387800_ref10","doi-asserted-by":"crossref","first-page":"902","DOI":"10.1038\/s41467-021-21229-w","article-title":"Multifunctional evolution of B and AGL6 MADS box genes in orchids","volume":"12","author":"Hsu","year":"2021","journal-title":"Nat Commun."},{"key":"2026021605165387800_ref11","doi-asserted-by":"crossref","first-page":"1328","DOI":"10.1093\/pcp\/pcae069","article-title":"PaWOX3 and PaWOX3B regulate flower number and the lip symmetry of Phalaenopsis","volume":"65","author":"Hsu","year":"2024","journal-title":"Plant Cell Physiol."},{"issue":"1","key":"2026021605165387800_ref12","doi-asserted-by":"crossref","first-page":"8","DOI":"10.2503\/hortj.MI-IR05","article-title":"Molecular mechanism regulating floral architecture in monocotyledonous ornamental Plants","volume":"85","author":"Kanno","year":"2016","journal-title":"Hortic. J."},{"key":"2026021605165387800_ref13","doi-asserted-by":"crossref","first-page":"831","DOI":"10.1023\/A:1025070827979","article-title":"Heterotopic expression of class B floral homeotic genes supports a modified ABC model for tulip (Tulipa gesneriana)","volume":"52","author":"Kanno","year":"2003","journal-title":"Plant Mol. Biol."},{"key":"2026021605165387800_ref14","doi-asserted-by":"crossref","first-page":"268","DOI":"10.1100\/tsw.2007.86","article-title":"Class B gene expression and the modified ABC model in nongrass monocots","volume":"7","author":"Kanno","year":"2007","journal-title":"ScientificWorldJournal"},{"key":"2026021605165387800_ref15","doi-asserted-by":"crossref","first-page":"750","DOI":"10.1105\/tpc.107.050385","article-title":"Elaboration of B gene function to include the identity of novel floral organs in the lower eudicot Aquilegia","volume":"19","author":"Kramer","year":"2007","journal-title":"Plant Cell"},{"key":"2026021605165387800_ref16","first-page":"pdb.prot4954","volume-title":"Phloroglucinol stain for lignin","author":"Liljegren","year":"2010"},{"key":"2026021605165387800_ref17","doi-asserted-by":"crossref","first-page":"129","DOI":"10.1016\/j.semcdb.2009.11.019","article-title":"The ABC model and the diversification of floral organ identity","volume":"21","author":"Litt","year":"2010","journal-title":"Semin. Cell Dev. Biol."},{"key":"2026021605165387800_ref18","doi-asserted-by":"crossref","first-page":"738","DOI":"10.1094\/MPMI-10-11-0266","article-title":"A high-throughput virus-induced gene-silencing vector for screening transcription factors in virus-induced plant defense response in orchid","volume":"25","author":"Lu","year":"2012","journal-title":"Mol. Plant Microbe Interact."},{"key":"2026021605165387800_ref19","first-page":"2079","article-title":"The C-terminal sequence and PI motif of the orchid (Oncidium Gower Ramsey) PI ortholog determines its ability to bind AP3 orthologs and enter the nucleus to regulate downstream genes controlling petal and stamen formation","volume":"56","author":"Mao","year":"2015","journal-title":"Plant Cell Physiol."},{"key":"2026021605165387800_ref20","doi-asserted-by":"crossref","first-page":"1357","DOI":"10.1111\/tpj.15117","article-title":"Distance-based measurement determines the coexistence of B protein hetero- and homodimers in lily tepal and stamen tetrameric complexes","volume":"105","author":"Mao","year":"2021","journal-title":"Plant J."},{"key":"2026021605165387800_ref21","doi-asserted-by":"crossref","first-page":"853","DOI":"10.1016\/j.tplants.2019.05.012","article-title":"Regulation of cell wall thickening by a medley of mechanisms","volume":"24","author":"McCahill","year":"2019","journal-title":"Trends Plant Sci."},{"key":"2026021605165387800_ref22","doi-asserted-by":"crossref","first-page":"831","DOI":"10.3389\/fpls.2018.00831","article-title":"The greenish flower phenotype of Habenaria radiata (Orchidaceae) is caused by a mutation in the SEPALLATA-Like MADS-Box gene HrSEP-1","volume":"9","author":"Mitoma","year":"2018","journal-title":"Front. Plant Sci."},{"key":"2026021605165387800_ref23","doi-asserted-by":"crossref","first-page":"51","DOI":"10.1016\/j.tplants.2007.11.007","article-title":"MADS about the evolution of orchid flowers","volume":"13","author":"Mondragon-Palomino","year":"2008","journal-title":"Trends Plant Sci."},{"key":"2026021605165387800_ref24","doi-asserted-by":"crossref","first-page":"1008","DOI":"10.1111\/j.1365-313X.2011.04560.x","article-title":"Conserved differential expression of paralogous DEFICIENS- and GLOBOSA-like MADS-box genes in the flowers of Orchidaceae: refining the \u2018orchid code\u2019","volume":"66","author":"Mondragon-Palomino","year":"2011","journal-title":"Plant J."},{"key":"2026021605165387800_ref25","doi-asserted-by":"crossref","first-page":"343","DOI":"10.1104\/pp.97.1.343","article-title":"Dihydroflavonol reductase activity in relation to differential anthocyanin accumulation in juvenile and mature phase Hedera helix L","volume":"97","author":"Murray","year":"1991","journal-title":"Plant Physiol."},{"key":"2026021605165387800_ref26","doi-asserted-by":"crossref","first-page":"288","DOI":"10.3389\/fpls.2015.00288","article-title":"NAC-MYB-based transcriptional regulation of secondary cell wall biosynthesis in land plants","volume":"6","author":"Nakano","year":"2015","journal-title":"Front. Plant Sci."},{"key":"2026021605165387800_ref27","doi-asserted-by":"crossref","first-page":"1024","DOI":"10.1111\/nph.12723","article-title":"Flower development of Phalaenopsis orchid involves functionally divergent SEPALLATA-like genes","volume":"202","author":"Pan","year":"2014","journal-title":"New Phytol."},{"key":"2026021605165387800_ref28","doi-asserted-by":"crossref","first-page":"345","DOI":"10.1093\/genetics\/140.1.345","article-title":"Molecular evolution of flower development: diversification of the plant MADS-box regulatory gene family","volume":"140","author":"Purugganan","year":"1995","journal-title":"Genetics"},{"key":"2026021605165387800_ref29","doi-asserted-by":"crossref","first-page":"399","DOI":"10.3389\/fpls.2018.00399","article-title":"Current models for transcriptional regulation of secondary cell wall biosynthesis in grasses","volume":"9","author":"Rao","year":"2018","journal-title":"Front Plant Sci."},{"key":"2026021605165387800_ref30","doi-asserted-by":"crossref","first-page":"96","DOI":"10.3732\/ajb.0800038","article-title":"One size fits all? Molecular evidence for a commonly inherited petal identity program in Ranunculales","volume":"96","author":"Rasmussen","year":"2009","journal-title":"Am J Bot."},{"key":"2026021605165387800_ref31","doi-asserted-by":"crossref","first-page":"1819","DOI":"10.1105\/tpc.106.042937","article-title":"Analysis of the Petunia TM6 MADS box gene reveals functional divergence within the DEF\/AP3 lineage","volume":"18","author":"Rijpkema","year":"2006","journal-title":"Plant Cell"},{"key":"2026021605165387800_ref32","first-page":"1259","article-title":"Diverse roles for MADS box genes in Arabidopsis development","volume":"7","author":"Rounsley","year":"1995","journal-title":"Plant Cell"},{"key":"2026021605165387800_ref33","doi-asserted-by":"crossref","first-page":"987","DOI":"10.1111\/j.1365-313X.2009.03928.x","article-title":"Intronic regulatory elements determine the divergent expression patterns of AGAMOUS-LIKE6 subfamily members in Arabidopsis","volume":"59","author":"Schauer","year":"2009","journal-title":"Plant J."},{"issue":"11","key":"2026021605165387800_ref34","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pone.0080462","article-title":"A modified ABCDE model of flowering in orchids based on gene expression profiling studies of the moth orchid Phalaenopsis aphrodite","volume":"8","author":"Su","year":"2013","journal-title":"PLoS One"},{"key":"2026021605165387800_ref35","doi-asserted-by":"crossref","first-page":"3022","DOI":"10.1093\/molbev\/msab120","article-title":"MEGA11: Molecular Evolutionary Genetics Analysis Version 11","volume":"38","author":"Tamura","year":"2021","journal-title":"Mol. Biol. Evol."},{"key":"2026021605165387800_ref36","doi-asserted-by":"crossref","first-page":"75","DOI":"10.1016\/S1369-5266(00)00139-4","article-title":"Development of floral organ identity: Stories from the MADS house","volume":"4","author":"Theissen","year":"2001","journal-title":"Curr. Opin. Plant Biol."},{"key":"2026021605165387800_ref37","doi-asserted-by":"crossref","first-page":"628","DOI":"10.1016\/0959-437X(95)80032-8","article-title":"MADS-box genes in ontogeny and phylogeny of plants: Haeckel's 'biogenetic law' revisited","volume":"5","author":"Theissen","year":"1995","journal-title":"Curr Opin Genet Dev"},{"key":"2026021605165387800_ref38","doi-asserted-by":"crossref","first-page":"469","DOI":"10.1038\/35054172","article-title":"Plant biology. Floral quartets","volume":"409","author":"Theissen","year":"2001","journal-title":"Nature"},{"key":"2026021605165387800_ref39","doi-asserted-by":"crossref","first-page":"115","DOI":"10.1023\/A:1006332105728","article-title":"A short history of MADS-box genes in plants","volume":"42","author":"Theissen","year":"2000","journal-title":"Plant Mol. Biol."},{"key":"2026021605165387800_ref40","doi-asserted-by":"crossref","first-page":"831","DOI":"10.1093\/pcp\/pch095","article-title":"Four DEF-like MADS box genes displayed distinct floral morphogenetic roles in Phalaenopsis orchid","volume":"45","author":"Tsai","year":"2004","journal-title":"Plant Cell Physiol."},{"key":"2026021605165387800_ref41","doi-asserted-by":"crossref","first-page":"1156","DOI":"10.1093\/pcp\/pce151","article-title":"A MADS box gene from lily (Lilium Longiflorum) is sufficient to generate dominant negative mutation by interacting with PISTILLATA (PI) in Arabidopsis thaliana","volume":"42","author":"Tzeng","year":"2001","journal-title":"Plant Cell Physiol."},{"key":"2026021605165387800_ref42","doi-asserted-by":"crossref","first-page":"741","DOI":"10.1105\/tpc.019166","article-title":"The duplicated B-class heterodimer model: whorl-specific effects and complex genetic interactions in Petunia hybrida flower development","volume":"16","author":"Vandenbussche","year":"2004","journal-title":"Plant Cell"},{"issue":"8","key":"2026021605165387800_ref43","doi-asserted-by":"crossref","first-page":"1065","DOI":"10.1111\/jipb.12891","article-title":"Split Nano luciferase complementation for probing protein-protein interactions in plant cells","volume":"62","author":"Wang","year":"2019","journal-title":"J. of Integrative Plant Biology"},{"key":"2026021605165387800_ref44","doi-asserted-by":"crossref","first-page":"353","DOI":"10.1111\/ppl.12647","article-title":"B and E MADS-box genes determine the perianth formation in Cymbidium goeringii Rchb.f","volume":"162","author":"Xiang","year":"2018","journal-title":"Physiol. Plant."},{"key":"2026021605165387800_ref45","doi-asserted-by":"crossref","first-page":"1923","DOI":"10.1100\/tsw.2006.320","article-title":"Function and diversification of MADS-box genes in rice","volume":"6","author":"Yamaguchi","year":"2006","journal-title":"TheScientificWorldJournal"},{"key":"2026021605165387800_ref46","doi-asserted-by":"crossref","first-page":"227","DOI":"10.1016\/j.tplants.2010.12.005","article-title":"Transcriptional networks for lignin biosynthesis: more complex than we thought?","volume":"16","author":"Zhao","year":"2011","journal-title":"Trends Plant Sci."},{"key":"2026021605165387800_ref47","doi-asserted-by":"crossref","first-page":"2763","DOI":"10.1105\/tpc.108.061325","article-title":"A battery of transcription factors involved in the regulation of secondary cell wall biosynthesis in Arabidopsis","volume":"20","author":"Zhong","year":"2008","journal-title":"Plant Cell"},{"key":"2026021605165387800_ref48","doi-asserted-by":"crossref","first-page":"248","DOI":"10.1105\/tpc.108.063321","article-title":"MYB58 and MYB63 are transcriptional activators of the lignin biosynthetic pathway during secondary cell wall formation in Arabidopsis","volume":"21","author":"Zhou","year":"2009","journal-title":"Plant Cell"},{"issue":"8","key":"2026021605165387800_ref49","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pone.0105726","article-title":"Arabidopsis NAC domain proteins, VND1 to VND5, are transcriptional regulators of secondary wall biosynthesis in vessels","volume":"9","author":"Zhou","year":"2014","journal-title":"PLoS ONE"}],"container-title":["Plant and Cell Physiology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/academic.oup.com\/pcp\/advance-article-pdf\/doi\/10.1093\/pcp\/pcaf137\/64956309\/pcaf137.pdf","content-type":"application\/pdf","content-version":"am","intended-application":"syndication"},{"URL":"https:\/\/academic.oup.com\/pcp\/article-pdf\/67\/1\/67\/64956309\/pcaf137.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"https:\/\/academic.oup.com\/pcp\/article-pdf\/67\/1\/67\/64956309\/pcaf137.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,2,16]],"date-time":"2026-02-16T10:17:01Z","timestamp":1771237021000},"score":1,"resource":{"primary":{"URL":"https:\/\/academic.oup.com\/pcp\/article\/67\/1\/67\/8303900"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2025,10,27]]},"references-count":49,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2025,10,27]]},"published-print":{"date-parts":[[2026,1,19]]}},"URL":"https:\/\/doi.org\/10.1093\/pcp\/pcaf137","relation":{},"ISSN":["0032-0781","1471-9053"],"issn-type":[{"value":"0032-0781","type":"print"},{"value":"1471-9053","type":"electronic"}],"subject":[],"published-other":{"date-parts":[[2026,1]]},"published":{"date-parts":[[2025,10,27]]}}}