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. 2020 Mar 25;11:1549. doi: 10.1038/s41467-020-15331-8

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© The Author(s) 2020

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. 8 — A model through which intracellular c-di-GMP interactions with the extension ATPase MshE, modulates extension and retraction activity of the MSHA pilus to facilitate near-surface motility patterns and V. cholerae surface attachment. (Step 1) MSHA elaborated on the cell surface (red) facilitate initial interactions of a motile cell (gray) with the surface. Black dots in the model are indicative of MSHA interactions with the surface. As motile cells have low levels of intracellular c-di-GMP, conditions are primed to promote MSHA PilT-mediated retraction via reduced MshE extension activity. At this stage, flagellar motion (green) can continue (indicated by black circle around the flagellum) resulting in near-surface orbiting motility. (Step 2) MSHA pilus retraction aids in facilitating initial surface interactions and attachment, and then yet to be characterized intracellular signaling cascade(s) initiate an increase in intracellular c-di-GMP production that results in attenuation of flagellar-mediated motion. (Step 3) Increasing intracellular c-di-GMP levels promote MshE-mediated extension activity, to allow for further MSHA production on the cell surface and enable anchoring of the cell to the surface. Cells which do not increase intracellular c-di-GMP levels upon initial surface interactions, demonstrate extended periods of PilT-mediated retraction that can promote transient interactions and detachment from the surface. (Step 4) Upon cell division, newly produced daughter cells (blue) can either remain attached to the surface to promote microcolony formation, or return to a planktonic free-swimming state. Prolonged interactions with the surface and continual increase of intracellular c-di-GMP levels will allow for alterations in gene expression dynamics for production of components necessary for biofilm formation. (Step 5) Continued production of extracellular matrix components (matrix), and cellular replication furthers biofilm formation and maturation.