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. 2019 Jun 10;9:8388. doi: 10.1038/s41598-019-44640-2

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© The Author(s) 2019

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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The proposed genetic clutch. (a) When a promoter-containing GC inserts into the first position, it can act as a genetic clutch by disengaging the original first GC (blue arrow) from P_C promoter and replaced with the one on promoter GC. When a new GC (green arrow) inserts, it can be expressed by P_C promoter, while the blue GC is expressed by promoter-containing GC and P_C promoter. (b) The expression level of gene cassettes with and without a genetic clutch. The estimated levels of expression of the blue ORF in (i) the first, (ii) the second and (iii) the third position were shown in the bar chart. The solid bars represent the situation when promoter-containing GC was inserted upstream of the blue GC, while the gridded bars represent the situation when no promoter-containing GC was inserted. The asterisks indicate the experimentally verified expression level, suggested by the results in Fig. 4 (TMB4 P_C and TMB4 P_C + GC). The expression of the blue ORF was hypothesised to be decreased when more GCs are inserted without the presence of a promoter-containing GC as a genetic clutch (gridded bars), based on the data from previous study⁴⁹.