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Substitution spectrum and selection at G-quadruplexes in great ape telomere-to-telomere genomes

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G-quadruplexes (G4s) are noncanonical DNA secondary structures formed by runs of guanines (stems) connected by other nucleotides (loops). These structures are enriched at regulatory regions such as promoters, CpG islands, untranslated regions (UTRs), enhancers... ...