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. 2020 Sep 23;10:15494. doi: 10.1038/s41598-020-72543-0

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© The Author(s) 2020

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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Model of the Tyr locus. (a) Representative individuals of mouse models for the Tyr locus: YRT2, TYRINS3, TYRINS5 and albino NMRI, along with (b) schematics of their genomic modifications. (c) When transcriptionally active, the Tyr locus is organized in two loops, mediated by CTCF at the 3′ boundary and likely by Sox10, Mitf and Usf1 binding at the 5′ boundary. (d) Inactivation of the Tyr 3′ boundary results in ectopic expression of Nox4, the gene located beyond the boundary. (e) Inactivation of the Tyr 5′ boundary causes loss of Tyr expression and overexpression of Grm5, past the boundary. Figure prepared with Adobe Illustrator. The image included in this figure has been obtained by Davide Seruggia at the CNB-CSIC in Madrid under the supervision of Lluis Montoliu.