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. 2017 Sep 8;7:11038. doi: 10.1038/s41598-017-10633-2

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© The Author(s) 2017

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 license, and indicate if changes were made. The images or other third party material in this article are included in the article’s Creative Commons license, unless indicated otherwise in a credit line to the material. If material is not included in the article’s Creative Commons license 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 license, visit http://creativecommons.org/licenses/by/4.0/.

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Dynamics of an imperfect suppression HD, where the drive fails and produces naturally resistant alleles, alone (a) and with countermeasures (b–g), which include a synthetic resistant allele (b,c), reversal drive (d,e), and immunizing reversal drive (f,g). Homing is imperfect (e _H = e _C = 0.9), and unsuccessful homing results in natural resistance via NHEJ with fitness cost s _R = 0.05. See Fig. 1 for other details. The biggest change from accounting for imperfect homing is that the IRD falls out of the population in the long-term (f,g) because of low-fitness cost alleles resistant to cutting.