Cortex cis-regulatory switches establish scale colour identity and pattern diversity in Heliconius

Luca Livraghi, Joseph J Hanly, Steven M Van Bellghem, Gabriela Montejo-Kovacevich, Eva Sm van der Heijden, Ling Sheng Loh, Anna Ren, Ian A Warren, James J Lewis, Carolina Concha, Laura Hebberecht, Charlotte J Wright, Jonah M Walker, Jessica Foley, Zachary H Goldberg, Henry Arenas-Castro, Camilo Salazar, Michael W Perry, Riccardo Papa, Arnaud MartinW Owen McMillan, Chris D Jiggins

Research output: Contribution to journalArticlepeer-review

30 Scopus citations


In Heliconius butterflies, wing colour pattern diversity and scale types are controlled by a few genes of large effect that regulate colour pattern switches between morphs and species across a large mimetic radiation. One of these genes, cortex, has been repeatedly associated with colour pattern evolution in butterflies. Here we carried out CRISPR knockouts in multiple Heliconius species and show that cortex is a major determinant of scale cell identity. Chromatin accessibility profiling and introgression scans identified cis-regulatory regions associated with discrete phenotypic switches. CRISPR perturbation of these regions in black hindwing genotypes recreated a yellow bar, revealing their spatially limited activity. In the H. melpomene/timareta lineage, the candidate CRE from yellow-barred phenotype morphs is interrupted by a transposable element, suggesting that cis-regulatory structural variation underlies these mimetic adaptations. Our work shows that cortex functionally controls scale colour fate and that its cis-regulatory regions control a phenotypic switch in a modular and pattern-specific fashion.

Original languageEnglish (US)
Article numbere68549
StatePublished - Jul 19 2021

All Science Journal Classification (ASJC) codes

  • General Neuroscience
  • General Biochemistry, Genetics and Molecular Biology
  • General Immunology and Microbiology


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