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. 2026 Sep 16;10:292. doi: 10.1038/s41538-026-01162-2

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© The Author(s) 2026

Open Access This article is licensed under a Creative Commons Attribution-NonCommercial-NoDerivatives 4.0 International License, which permits any non-commercial use, sharing, 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 you modified the licensed material. You do not have permission under this licence to share adapted material derived from this article or parts of it. 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-nc-nd/4.0/.

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Fig. 6 — A Map of pathways associated with TGF-β/Smad and Bmp4/Smad signaling (Created with BioGDP.com). B Heat map of mRNA expression levels of gene TGF-β2, TGF-β3, Smad2, Smad3, Smad4, Serpina5, Bmp4, Bmpr1α, Bmpr2, Bmpr1β, Smad1, Smad5, FGF2 and Kitl in testicular tissue (n = 5). C, D Immunofluorescence staining of Bmp4 (red) and Bmpr1α (red) protein in testicular tissue (Scale bars = 50 μm, n = 5). Nuclei were counterstained with DAPI (blue). E–H Molecular docking results of Bmp4-Riboflavin, Bmpr1α-Riboflavin, Smad4-Riboflavin, and Smad5-Riboflavin, respectively. Note: Control: Control group; As: Arsenic trioxide group; Ribo: riboflavin group; As + Ribo: co-administration of arsenic trioxide and riboflavin group. ***P < 0.001, **P < 0.01 and *P < 0.05 vs. Control group; ^###P < 0.001, ^##P < 0.01 and ^#P < 0.05 vs. As group.