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. 2022 Nov 8;5:146. doi: 10.1038/s42004-022-00765-6

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© The Author(s) 2022

Open Access This article is licensed under a Creative Commons Attribution 4.0 International License, which permits use, sharing, adaptation, distribution and reproduction in any medium or format, as long as you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons license, and indicate if changes were made. The images or other third party material in this article are included in the article’s Creative Commons license, unless indicated otherwise in a credit line to the material. If material is not included in the article’s Creative Commons license and your intended use is not permitted by statutory regulation or exceeds the permitted use, you will need to obtain permission directly from the copyright holder. To view a copy of this license, visit http://creativecommons.org/licenses/by/4.0/.

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Fig. 1 — a Structures of labdane (red), halimane (green), and clerodane (blue) skeletons. b The tpn BGC consists of two TSs (blue), two P450s (red), a ferredoxin (brown) and a GGPP synthase (green); mevalonate pathway genes responsible for production of the terpene precursors are found downstream. c Tpn2 first cyclizes GGPP into terpentedienyl diphosphate (2), a clerodane diterpene. Tpn3 eliminates the diphosphate of 2 to produce terpentetriene (3). The oxidation of 3 to terpentecin is still unknown. Carbon numbering for both 1 and 2 are shown. d HPLC analysis of the Tpn2 in vitro reaction confirms its type II TS activity. Negative control is reaction with boiled Tpn2. Products are shown as the dephosphorylated analogs.