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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 detection of the integron GCs by using pBiDiPD. (a) pBiDiPD Transformants on LB agar supplemented with chloramphenicol, X-gal/IPTG and 4-methylumbelliferyl β-D-glucuronide (MUG), (b) Blue-white screening to detect for the clones with promoter activity on the antisense strand, (c) Exposing the colonies under the UV light to detect clones with promoter activity on the sense strand. The positive (+) and negative (−) colonies were the E. coli containing pCC1BAC-lacZα-TMB4-P_C-gusA (with experimentally proven promoter activities on either strand of DNA and pCC1BAC-lacZα-gusA (no promoter activity), respectively.