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. 2022 Apr 4;12:5606. doi: 10.1038/s41598-022-08998-0

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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 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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Seven-step approach to DNA methylation signature discovery, incorporating twin design. The figure represents the methodology of DNA methylation signature discovery study of a phenotype in multiple steps. It integrates behavioral-genetic and SNP-based methods (step 1) to estimate heritability, epigenome-wide study methods (steps 3–4) for association analyses, follow-up of results using summary statistics from previous EWASs and GWASs (step 5), the discordant twin design (step 6), and methods integrating polygenic and DNA methylation data (step 7) in enrichment analysis. Specific methods for each step are presented on the left and outcomes on the right. GWAS, genome-wide association study. EWAS, epigenome-wide association study. SNP, single nucleotide polymorphisms. CpG, cytosine-phosphate-guanine. PGS, polygenic scores. MS, methylation scores. DMPs, differentially methylated positions. DMRs, differentially methylated regions.