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. 2018 Aug 1;8:11552. doi: 10.1038/s41598-018-29884-8

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© The Author(s) 2018

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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Physiology of flagellin/FliS binding to FlhA-C. (a) Projection of different amino acid exchanges onto the structure of flagellin/FliS. Flagellin is shown in orange, while FliS is depicted in blue. Dashed lines indicate disordered regions. (b–e) Quantitative swarm expansion assay for fliS and flagellin (hag) mutants: WT is swarming proficient and covers a 0.7% LB agar plate in about 4.5 hours but ΔfliS is non-swarming. Both fliS^Δ2–18 and fliS^Y7A,Y10A phenocopy a ΔfliS mutant for swarming. fliS^K33E is swarming proficient. hag^R57E confers a non-swarming phenotype while all hag mutants except hag^V299D show almost no swarming motility. (f–o) Fluorescence microscopy of B. subtilis filaments shows deficiency of ΔfliS, fliS^Δ2-18 and fliS^Y7A,Y10A mutants to secrete flagellin. fliS^K33E shows short flagellar filaments. hag^R57E, hag^V299D, hag^L300E and hag^R304E show almost no flagellar filaments, while hag^Q297A exhibits short flagellar filaments. Scale bars are 2 µm.