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Significance C:G to T:A mutations constitute the largest class of spontaneous base substitutions in all organisms. These mutations are thought to be a result of cytosine deaminations, but what promotes these deaminations is unclear. We confirm here the hypothesis that they occur predominantly in single-stranded DNA (ssDNA) and identify the ssDNA in the lagging strand template as the preferred site of C:G to T:A mutations. As a consequence, replication creates a strand bias in these mutations, and this overwhelms any strand bias resulting from transcription. These results explain a long-recognized bias in base composition of microbial genomes called GC skew and predicts that C:G to T:A mutations created by the APOBEC3 family deaminases in cancer genomes should occur with the same strand bias.

Strand-biased cytosine deamination at the replication fork causes cytosine to thymine mutations in Escherichia coli