{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,2,4]],"date-time":"2026-02-04T19:10:36Z","timestamp":1770232236007,"version":"3.49.0"},"reference-count":43,"publisher":"Frontiers Media SA","license":[{"start":{"date-parts":[[2026,2,4]],"date-time":"2026-02-04T00:00:00Z","timestamp":1770163200000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100008402","name":"Chengdu University of Traditional Chinese Medicine","doi-asserted-by":"publisher","award":["030040015"],"award-info":[{"award-number":["030040015"]}],"id":[{"id":"10.13039\/501100008402","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["frontiersin.org"],"crossmark-restriction":true},"short-container-title":["Front. Plant Sci."],"abstract":"<jats:sec>\n                    <jats:title>Introduction<\/jats:title>\n                    <jats:p>\n                      The opium poppy (\n                      <jats:italic>Papaver somniferum<\/jats:italic>\n                      L.) is the primary source of medically important benzylisoquinoline alkaloids (BIAs), including morphine and codeine. Nevertheless, the transcriptional regulatory networks and transport processes underlying the spatiotemporal accumulation of BIAs during capsule development remain incompletely understood.\n                    <\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Methods<\/jats:title>\n                    <jats:p>We performed an integrated transcriptomic and metabolomic analysis across five defined capsule developmental stages (S1\u2013S5). Transcript\u2013metabolite relationships were examined using global correlation analysis, trend analysis, and weighted gene co-expression network analysis (WGCNA). To identify putative BIA transporters, we applied a multi-tiered bioinformatic screening pipeline that combined hub-gene prioritization, transmembrane domain prediction, and functional annotation, followed by qRT-PCR validation of shortlisted candidates.<\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Results<\/jats:title>\n                    <jats:p>\n                      Metabolomic profiling revealed a clear developmental trajectory of BIA accumulation, with S4 emerging as a critical stage for BIA biosynthesis. Terminal alkaloids reached maximal levels at S4 and declined sharply at S5, suggesting the initiation of active transport and\/or metabolic conversion at late development. Strong transcript\u2013metabolite concordance enabled robust multi-omics integration. WGCNA identified 32 co-expression modules, among which the steelblue, brown, blue, and white modules showed the strongest associations with BIA metabolites, including the key intermediate thebaine. The integrated screening strategy and qRT-PCR validation ultimately highlighted\n                      <jats:italic>PsMATE1<\/jats:italic>\n                      and\n                      <jats:italic>PsEXS1<\/jats:italic>\n                      as the highest-confidence transporter candidates, supported by their multiple predicted transmembrane domains, membership in established transporter families, and expression patterns tightly synchronized with late-stage BIA accumulation.\n                    <\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Conclusion<\/jats:title>\n                    <jats:p>\n                      This study provides a comprehensive multi-omics resource and a systematic framework for transporter discovery in P. somniferum capsules. The identification of\n                      <jats:italic>PsMATE1<\/jats:italic>\n                      and\n                      <jats:italic>PsEXS1<\/jats:italic>\n                      offers prioritized targets for future functional characterization and advances understanding of the mechanisms controlling BIA transport and accumulation during capsule development.\n                    <\/jats:p>\n                  <\/jats:sec>","DOI":"10.3389\/fpls.2026.1754793","type":"journal-article","created":{"date-parts":[[2026,2,4]],"date-time":"2026-02-04T06:47:54Z","timestamp":1770187674000},"update-policy":"https:\/\/doi.org\/10.3389\/crossmark-policy","source":"Crossref","is-referenced-by-count":0,"title":["An integrated transcriptomic and metabolomic atlas reveals the temporal regulation of benzylisoquinoline alkaloid biosynthesis and transport in developing opium poppy capsules"],"prefix":"10.3389","volume":"17","author":[{"given":"Qian","family":"Wang","sequence":"first","affiliation":[{"name":"School of Pharmacy, Chengdu University of Traditional Chinese Medicine","place":["Chengdu, China"]},{"name":"Institute of Herbgenomics, Chengdu University of Traditional Chinese Medicine","place":["Chengdu, China"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Xiaofang","family":"Qie","sequence":"additional","affiliation":[{"name":"School of Pharmacy, Chengdu University of Traditional Chinese Medicine","place":["Chengdu, China"]},{"name":"Institute of Herbgenomics, Chengdu University of Traditional Chinese Medicine","place":["Chengdu, China"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Yun","family":"Zhen","sequence":"additional","affiliation":[{"name":"School of Pharmacy, Chengdu University of Traditional Chinese Medicine","place":["Chengdu, China"]},{"name":"Institute of Herbgenomics, Chengdu University of Traditional Chinese Medicine","place":["Chengdu, China"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Ruoshi","family":"Li","sequence":"additional","affiliation":[{"name":"School of Pharmacy, Chengdu University of Traditional Chinese Medicine","place":["Chengdu, China"]},{"name":"Institute of Herbgenomics, Chengdu University of Traditional Chinese Medicine","place":["Chengdu, China"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Zhaoyu","family":"Liu","sequence":"additional","affiliation":[{"name":"College of Tropical Crops and College of Forestry, Hainan University","place":["Haikou, China"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Yanjun","family":"Zhang","sequence":"additional","affiliation":[{"name":"Wuhan Botanical Garden, Chinese Academy of Sciences","place":["Wuhan, China"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Shilin","family":"Chen","sequence":"additional","affiliation":[{"name":"School of Pharmacy, Chengdu University of Traditional Chinese Medicine","place":["Chengdu, China"]},{"name":"Institute of Herbgenomics, Chengdu University of Traditional Chinese Medicine","place":["Chengdu, China"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Chi","family":"Song","sequence":"additional","affiliation":[{"name":"School of Pharmacy, Chengdu University of Traditional Chinese Medicine","place":["Chengdu, China"]},{"name":"Institute of Herbgenomics, Chengdu University of Traditional Chinese Medicine","place":["Chengdu, China"]}],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"1965","published-online":{"date-parts":[[2026,2,4]]},"reference":[{"key":"B1","doi-asserted-by":"publisher","first-page":"700","DOI":"10.1186\/s12870-024-05412-x","article-title":"Developing benzylisoquinoline alkaloid-enriched opium poppy via CRISPR-directed genome editing: A review","volume":"24","author":"Aghaali","year":"2024","journal-title":"BMC Plant Biol."},{"key":"B2","doi-asserted-by":"publisher","first-page":"5941","DOI":"10.1128\/JB.00666-12","article-title":"Small multidrug resistance protein emrE reduces host pH and osmotic tolerance to metabolic quaternary cation osmoprotectants","volume":"194","author":"Bay","year":"2012","journal-title":"J. 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