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. 2021 Jul 5;12:4132. doi: 10.1038/s41467-021-23889-0

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© The Author(s) 2021

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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Fig. 1 — a Expression from promoters with no control circuitry (i.e. open-loop circuits) is proportional to the plasmid copy number. b An ideal dosage compensation system uses control mechanisms to tune the per-copy expression rate, thereby maintaining constant expression regardless of copy number.