{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,15]],"date-time":"2026-09-15T15:52:24Z","timestamp":1789487544155,"version":"build-2803163510"},"reference-count":115,"publisher":"American Chemical Society (ACS)","issue":"36","license":[{"start":{"date-parts":[[2026,9,2]],"date-time":"2026-09-02T00:00:00Z","timestamp":1788307200000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100001809","name":"National Natural Science Foundation of China","doi-asserted-by":"publisher","award":["22508155"],"award-info":[{"award-number":["22508155"]}],"id":[{"id":"10.13039\/501100001809","id-type":"DOI","asserted-by":"publisher"}]},{"name":"Key Project at Central Government Level: The Ability Establishment of Sustainable Use for Valuable Chinese Medicine Resources","award":["2060302"],"award-info":[{"award-number":["2060302"]}]},{"name":"Major Science and Technology Program of Yunnan Province","award":["202102AA3100454-5"],"award-info":[{"award-number":["202102AA3100454-5"]}]},{"name":"Yunnan Fundamental Research","award":["202501CF070130"],"award-info":[{"award-number":["202501CF070130"]}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2026,9,15]]},"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>Protopanaxadiol (PPD)-type ginsenosides, as the primary bioactive components of Panax medicinal plants, have become a research hotspot in natural products due to their diverse pharmacological activities. The biosynthetic pathway of PPD-type ginsenosides in Panax species and the related enzymes is becoming increasingly clear. However, for most enzymes, research has been largely limited to identifying their functions in the pathway, with insufficient exploration of their biochemical properties, structures, and mechanisms. This review, from the perspective of enzymatic catalysis, systematically elucidates the biosynthetic pathway of PPD-type ginsenosides in Panax species. It focuses on the functions, sequence, and biochemical features of the enzymes and uses homology modeling and AlphaFold predictions to generate testable hypotheses regarding their structures and catalytic mechanisms, while also summarizing directed evolution strategies, aiming to provide a theoretical foundation for the construction of in vitro or microbial \u201ccell factories\u201d and to promote the efficient synthesis and application of highly bioactive PPD-type ginsenosides.<\/jats:p>","DOI":"10.1021\/acsomega.6c06948","type":"journal-article","created":{"date-parts":[[2026,9,3]],"date-time":"2026-09-03T17:05:23Z","timestamp":1788455123000},"page":"53661-53679","source":"Crossref","is-referenced-by-count":0,"title":["Insights into Enzymes\nInvolved in the Biosynthesis\nof Protopanaxadiol-Type Ginsenosides in\n                    <i>Panax<\/i>\n                    Species"],"prefix":"10.1021","volume":"11","author":[{"ORCID":"https:\/\/orcid.org\/0000-0003-1488-8816","authenticated-orcid":true,"given":"Na","family":"Li","sequence":"first","affiliation":[{"name":"Kunming University of Science and Technology , , ,","place":["Kunming, China, 650500"]},{"name":"Key Laboratory of Sustainable Utilization of Panax Notoginseng Resources of Yunnan Province , ,","place":["Kunming, China, 650500"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xingyi","family":"Jiang","sequence":"additional","affiliation":[{"name":"Kunming University of Science and Technology , , ,","place":["Kunming, China, 650500"]},{"name":"Key Laboratory of Sustainable Utilization of Panax Notoginseng Resources of Yunnan Province , ,","place":["Kunming, China, 650500"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0002-7163-3641","authenticated-orcid":true,"given":"Shisong","family":"Gao","sequence":"additional","affiliation":[{"name":"Kunming University of Science and Technology , , ,","place":["Kunming, China, 650500"]},{"name":"Key Laboratory of Sustainable Utilization of Panax Notoginseng Resources of Yunnan Province , ,","place":["Kunming, China, 650500"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Qin","family":"Chen","sequence":"additional","affiliation":[{"name":"Kunming University of Science and Technology , , ,","place":["Kunming, China, 650500"]},{"name":"Key Laboratory of Sustainable Utilization of Panax Notoginseng Resources of Yunnan Province , ,","place":["Kunming, China, 650500"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-1127-7120","authenticated-orcid":true,"given":"Feng","family":"Ge","sequence":"additional","affiliation":[{"name":"Kunming University of Science and Technology , , ,","place":["Kunming, China, 650500"]},{"name":"Key Laboratory of Sustainable Utilization of Panax Notoginseng Resources of Yunnan Province , ,","place":["Kunming, China, 650500"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xiuming","family":"Cui","sequence":"additional","affiliation":[{"name":"Kunming University of Science and Technology , , ,","place":["Kunming, China, 650500"]},{"name":"Key Laboratory of Sustainable Utilization of Panax Notoginseng Resources of Yunnan Province , ,","place":["Kunming, China, 650500"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"316","published-online":{"date-parts":[[2026,9,2]]},"reference":[{"issue":"7","key":"2026091510091423400_cit1","doi-asserted-by":"publisher","first-page":"1813","DOI":"10.1016\/j.apsb.2020.12.017","article-title":"Ginsenosides in Panax Genus and Their Biosynthesis","volume":"11","author":"Hou","year":"2021","journal-title":"Acta Pharm. 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Wellness"},{"issue":"5","key":"2026091510091423400_cit5","doi-asserted-by":"publisher","first-page":"803","DOI":"10.1016\/j.jfutfo.2024.11.016","article-title":"Panax notoginseng Saponins Attenuate Oxidative Stress-Mediated\nPlatelet Apoptosis\nin Hyperlipidemic Conditions via Up-Regulating cAMP\/PKA Signaling\nPathway In Vitro and In Vivo","volume":"6","author":"Ma","year":"2026","journal-title":"J. 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Plants"},{"key":"2026091510091423400_cit29","first-page":"611","article-title":"Effect of Farnesyl\nPyrophosphate Synthase (FPS) on Saponin Biosynthesis\nin Panax japonicus","volume":"38","author":"Liu","year":"2018","journal-title":"Bull. Bot.\nRes."},{"issue":"18","key":"2026091510091423400_cit30","doi-asserted-by":"publisher","first-page":"2821","DOI":"10.3390\/plants14182821","article-title":"Sustainable\nProduction of Ginsenosides: Advances in Biosynthesis and Metabolic\nEngineering","volume":"14","author":"Xue","year":"2025","journal-title":"Plants (Basel)"},{"key":"2026091510091423400_cit31","doi-asserted-by":"publisher","first-page":"658","DOI":"10.1016\/j.ijbiomac.2016.11.070","article-title":"Molecular Cloning and Functional Analysis of Squalene Synthase (SS)\nin Panax notoginseng","volume":"95","author":"Jiang","year":"2017","journal-title":"Int.\nJ. Biol. Macromol."},{"key":"2026091510091423400_cit32","doi-asserted-by":"publisher","first-page":"38","DOI":"10.1016\/j.bej.2017.03.001","article-title":"Enhancement\nof Triterpenoid Saponins Biosynthesis in Panax notoginseng Cells by Co-overexpressions of 3-hydroxy-3-methylglutaryl CoA Reductase\nand Squalene Synthase Genes","volume":"122","author":"Deng","year":"2017","journal-title":"Biochem. Eng. J."},{"key":"2026091510091423400_cit33","doi-asserted-by":"publisher","first-page":"182","DOI":"10.1016\/j.indcrop.2018.02.036","article-title":"Methyl Jasmonate as\na Control Factor of the Synthase\nSqualene Gene Promoter and Ginsenoside Production in American Ginseng\nHairy Root Cultured in Shake Flasks and a Nutrient Sprinkle Bioreactor","volume":"115","author":"Kochan","year":"2018","journal-title":"Ind. 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Biotechnol."},{"issue":"5","key":"2026091510091423400_cit46","doi-asserted-by":"publisher","first-page":"506","DOI":"10.3390\/horticulturae10050506","article-title":"Effect of Phosphate-Deficiency\nStress on the Biological Characteristics and Transcriptomics of Panax ginseng","volume":"10","author":"Sun","year":"2024","journal-title":"Horticulturae"},{"issue":"8","key":"2026091510091423400_cit47","doi-asserted-by":"publisher","first-page":"614","DOI":"10.1016\/S1875-5364(22)60198-5","article-title":"Integrative\nSMRT Sequencing and Ginsenoside Profiling Analysis Provide Insights\ninto the Biosynthesis of Ginsenoside in Panax quinquefolium","volume":"20","author":"Di","year":"2022","journal-title":"Chin. J. Nat. 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