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. 2023 Jan 27;14:437. doi: 10.1038/s41467-023-36007-z

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© The Author(s) 2023

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 Semi-synthesis of 24-epi-brassinolide (EBL), 28-homo-brassinolide (HBL), and brassinolide (BL) from ergosterol, stigmasterol, and crinosterol, respectively. In comparison with crinosterol, 24-epi-ergosterol, the diastereoisomer to ergosterol, could be produced on a large scale using yeast strains constructed in the present study. b Manipulation of sterol homeostasis for the production of 24-epi-ergosterol in yeast. The introduction of a Δ²⁴⁽²⁸⁾-sterol reductase (DWF1) from plants enabled de novo biosynthesis of 24-epi-ergosterol. Afterward, the catalytic activity of DWF1 and, accordingly, the production of 24-epi-ergosterol was enhanced by directed evolution. The sterol fluxes towards 24-epi-ergosterol were further strengthened by the engineering of sterol homeostasis, maintaining a balance between sterol acylation (for storage in LDs) and steryl ester hydrolysis (for releasing free sterols in cellular membranes) via overexpression of YEH1, YEH2, and ARE2 with intact ARE1. DWF1 Δ²⁴⁽²⁸⁾-sterol reductase, ARE1 sterol O-acyltransferase 1, ARE2 sterol O-acyltransferase 2, YEH1 yeast steryl ester hydrolase 1, YEH2 yeast steryl ester hydrolase 2, TGL1 triglyceride lipase.