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. 2023 Nov 15;13:19927. doi: 10.1038/s41598-023-43099-6

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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 licence, and indicate if changes were made. The images or other third party material in this article are included in the article's Creative Commons licence, unless indicated otherwise in a credit line to the material. If material is not included in the article's Creative Commons licence 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 licence, visit http://creativecommons.org/licenses/by/4.0/.

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This scheme illustrates the proposed role of RAS in the heart of BO-treated mice. BO-treatment induces dual phosphorylation on PKCδ (Tyr-311/Thr-505), leading to CaMKII activation with the increased ROS production via activation of NOX4/XO and the ASK1-p38 MAPK cascade, which mediates the phosphorylation of RyR2 (Ser-2814) and PLB (Thr-17). BO-treatment was previously reported to induce sympathetic nerve activity by our group, leading to the phosphorylation of RyR2 (Ser-2808) and PLB (Ser-16)¹⁸. These changes might cause Ca²⁺ leakage, leading to fibrosis, myocyte apoptosis, oxidative stress and cardiac dysfunction. Solid black lines represent findings in this study and solid gray lines represents finding reported previously^18,20. β-AR, β-adrenoreceptor; Gsα, α subunit of Gs; AC5, type 5 adenylyl cyclase; SERCA, sarco/endoplastic reticulumn calcium ATPase; SR, sarcoplasmic reticulumn; RyR2, ryanodine receptor 2.