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. 2026 Feb 20;16:7463. doi: 10.1038/s41598-026-40642-z

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© The Author(s) 2026

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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Fig. 4 — Off-target mutation analysis of ABE-SpRY constructs. (a–d) Evaluation of cumulative on- and off-target DNA editing by ABE8e- and ABE9-SpRY in mouse embryos in a Cas9-dependent manner. Cumulative editing results from the sum of all A-to-G editing efficiencies within the spacer sequence. Control, gDNA from ear biopsy samples of adult wild-type mice. Individual data points for three independent biological replicates are shown. Error bars indicate standard deviation (SD); bounds were capped at 0% and 100% where the mean ± SD exceeded the valid range. Mismatches in the spacer sequence between on- and off-target sites are displayed in lowercase and magenta. (e) Diagram of the R-loop assay illustrating how the base editor can target transient non-target R-loops formed by dSaCas9. (f) Cas9-independent assessment of off-target DNA editing by ABE8e-SpRY and ABE9-SpRY, using the orthogonal R-loop assay at specific R-loop sites with five or six biological replicates. The black horizontal bar shows the mean at each adenine position grouped by editor. Parts of the schematics in (e) were created with BioRender.com.