{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,23]],"date-time":"2026-09-23T07:01:46Z","timestamp":1790146906066,"version":"4.0.1"},"reference-count":69,"publisher":"Frontiers Media SA","license":[{"start":{"date-parts":[[2026,9,23]],"date-time":"2026-09-23T00:00:00Z","timestamp":1790121600000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":["frontiersin.org"],"crossmark-restriction":true},"short-container-title":["Front. Plant Sci."],"abstract":"<jats:p>\n                    <jats:italic>Phlomoides rotata<\/jats:italic>\n                    , a Qinghai\u2013Tibet Plateau endemic perennial valued for its hemostatic and anti-inflammatory effects, derives part of these bioactivities from terpenoid secondary metabolism. While AP2\/ERF and bHLH transcription factors are established regulators of plant terpenoid biosynthesis, their roles in mediating\n                    <jats:italic>P. rotata<\/jats:italic>\n                    \u2019s adaptation to natural high-altitude gradients remain uncharacterized. Here, we integrated transcriptomic and metabolomic profiles from four wild\n                    <jats:italic>P. rotata<\/jats:italic>\n                    populations spanning the 3,540-4,270 m altitudinal gradient to investigate the associations of AP2\/ERF and bHLH with terpenoid biosynthesis. Genome-wide identification retrieved 169 AP2\/ERF and 212 bHLH family members, of which 71 AP2\/ERF and 50 bHLH genes displayed significant differential expression across four populations. Phylogenetic analysis resolved AP2\/ERF members into 15 subfamilies and bHLH members into 21 subfamilies, including a\n                    <jats:italic>P. rotata<\/jats:italic>\n                    -specific clade (subgroup XXII). We further focused on differential expression of AP2\/ERF and bHLH. Bioinformatic profiling revealed most encoded proteins are nuclear-localized, acidic, and unstable, with phytohormone-responsive cis-elements dominating AP2\/ERF promoters (39.31%) and abiotic stress-responsive elements enriched in bHLH promoters (41.55%). Weighted gene co-expression network analysis (WGCNA) identified eight modules, with the green module showing the strongest correlation with terpenoid metabolites (12 of 52 significant associations, |\n                    <jats:italic>r<\/jats:italic>\n                    | &amp;gt; 0.7,\n                    <jats:italic>p<\/jats:italic>\n                    \u00a0&amp;lt;\u00a00.01). Intra-modular analysis revealed distinct correlation patterns: specific AP2\/ERF and bHLH TFs exhibited strong positive correlations with iridoid glycosides, such as shanzhiside methyl ester, and key structural genes, including\n                    <jats:italic>IO<\/jats:italic>\n                    and\n                    <jats:italic>GGR<\/jats:italic>\n                    , but strong negative correlations with triterpenoid saponins, such as azukisaponin IV. qRT-PCR validation confirmed the reliability of the transcriptomic profiles. This study offers insights into the correlative framework linking AP2\/ERF and bHLH transcription factors with terpenoid secondary metabolism in\n                    <jats:italic>P. rotata<\/jats:italic>\n                    .\n                  <\/jats:p>","DOI":"10.3389\/fpls.2026.1957015","type":"journal-article","created":{"date-parts":[[2026,9,23]],"date-time":"2026-09-23T06:19:50Z","timestamp":1790144390000},"update-policy":"https:\/\/doi.org\/10.3389\/crossmark-policy","source":"Crossref","is-referenced-by-count":0,"title":["Characterization of AP2\/ERF and bHLH transcription factors and association with terpenoid metabolism in Phlomoides rotata from the Qinghai-Tibet Plateau"],"prefix":"10.3389","volume":"17","author":[{"given":"Rui","family":"Su","sequence":"first","affiliation":[{"name":"Key Laboratory of Tibetan Plateau Medicinal Plant and Animal Resources, School of Life Sciences, Qinghai Normal University","place":["Xining, Qinghai, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xiaoli","family":"Yang","sequence":"additional","affiliation":[{"name":"College of Ecology and Environmental Science, Qinghai Institute of Technology","place":["Xining, Qinghai, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yinglian","family":"Qi","sequence":"additional","affiliation":[{"name":"Key Laboratory of Tibetan Plateau Medicinal Plant and Animal Resources, School of Life Sciences, Qinghai Normal University","place":["Xining, Qinghai, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Guigong","family":"Geng","sequence":"additional","affiliation":[{"name":"Academy of Agricultural and Forestry Sciences, Qinghai University","place":["Xining, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Zirui","family":"Wang","sequence":"additional","affiliation":[{"name":"Key Laboratory of Tibetan Plateau Medicinal Plant and Animal Resources, School of Life Sciences, Qinghai Normal University","place":["Xining, Qinghai, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yaqi","family":"Li","sequence":"additional","affiliation":[{"name":"Key Laboratory of Tibetan Plateau Medicinal Plant and Animal Resources, School of Life Sciences, Qinghai Normal University","place":["Xining, Qinghai, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Feng","family":"Qiao","sequence":"additional","affiliation":[{"name":"Key Laboratory of Tibetan Plateau Medicinal Plant and Animal Resources, School of Life Sciences, Qinghai Normal University","place":["Xining, Qinghai, China"]},{"name":"Academy of Agricultural and Forestry Sciences, Qinghai University","place":["Xining, China"]},{"name":"Academy of Plateau Science and Sustainability, Qinghai Normal University","place":["Xining, Qinghai, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1965","published-online":{"date-parts":[[2026,9,23]]},"reference":[{"key":"B1","doi-asserted-by":"publisher","DOI":"10.3389\/fpls.2025.1718585","article-title":"Genome-wide identification and expression profiling of bHLH transcription factors associated with ferulic acid biosynthesis in angelica sinensis","volume":"16","author":"Arshad","year":"2025","journal-title":"Front. 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