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Here, we combine genomics, gene co-expression, and enzyme characterization to discover convergent 20E biosynthetic pathways from spinach and quinoa (amaranth family) and\n                  <jats:italic>Ajuga<\/jats:italic>\n                  species (mint family). We reveal that these families use distinct biosynthetic gene clusters to produce 20E and related phytoecdysteroids, highlighting both biochemical and genetic convergence of 20E biosynthesis in plants. Our work establishes the first complete 20E pathways in any organism and provides foundational insight into the evolution of this animal hormone across kingdoms of life.\n                <\/jats:p>","DOI":"10.64898\/2026.08.07.743582","type":"posted-content","created":{"date-parts":[[2026,8,10]],"date-time":"2026-08-10T13:35:19Z","timestamp":1786368919000},"source":"Crossref","is-referenced-by-count":0,"title":["Convergent biosynthesis of insect molting hormones in plants"],"prefix":"10.64898","author":[{"ORCID":"https:\/\/orcid.org\/0009-0008-1054-9212","authenticated-orcid":false,"given":"Joshua J.","family":"Park","sequence":"first","affiliation":[{"name":"Department of Molecular and Cellular Biology, Harvard University, Cambridge, MA, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-7831-534X","authenticated-orcid":false,"given":"Juan A.","family":"Orozco","sequence":"additional","affiliation":[{"name":"Department of Molecular and Cellular Biology, Harvard University, Cambridge, MA, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0122-5274","authenticated-orcid":false,"given":"Galen J. 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BMC Biotechnol. 8, 83 (2008).","DOI":"10.1186\/1472-6750-8-83"},{"key":"2026081121551928000_2026.08.07.743582v1.75","doi-asserted-by":"publisher","DOI":"10.1126\/science.1248000"},{"key":"2026081121551928000_2026.08.07.743582v1.76","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.2102949118"},{"key":"2026081121551928000_2026.08.07.743582v1.77","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-022-31100-1"},{"key":"2026081121551928000_2026.08.07.743582v1.78","doi-asserted-by":"crossref","first-page":"15326","DOI":"10.1021\/ja508127q","article-title":"Complete biosynthetic pathway of anditomin: Nature\u2019s sophisticated synthetic route to a complex fungal meroterpenoid","volume":"136","year":"2014","journal-title":"J. Am. Chem. 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