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. 2022 Jun 30;13:3780. doi: 10.1038/s41467-022-31587-8

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© The Author(s) 2022

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. 5 — a Claudin proteins expressed in nonepithelial cells form polymers similar to those within epithelial tight junction strands. b EGFP-claudin-2 expressed in U2OS human osteosarcoma cells forms strands (arrowhead). In contrast, mCherry-claudin-4 (human), claudin-4-EGFP (CLDN4-EGFP.C), untagged human claudin-4, canine claudin-4, and mouse claudin-4 all fail to form strands despite being trafficked to plasma membranes. Maximum projection images are representative of 3 independent experiments. Scale: 2 μm. c Live imaging at intervals after induction of mCherry-claudin-4 (magenta) expression shows progressive disruption of EGFP-claudin-2 (cyan) strands. Maximum projection images representative of 5 independent experiments. Scale: 10 μm; 2 μm. d Confocal and corresponding STED image of EGFP-claudin-2 polymers (cyan) without claudin-4 expression. STED shows collapse of EGFP-claudin-2 (cyan) polymers into dense aggregates (arrowhead) following mCherry-claudin-4 (magenta) expression (bottom). Representative of at least 7 independent experiments. Scale: 1 µm. e Quantitative morphometry shows progressive loss of EGFP-claudin-2 strands as mCherry-claudin-4 expression increases. n = 4 (each point is the mean of 12–16 cells in 1 sample), representative of 6 independent experiments. 1-way ANOVA. **P = 0.0035. f EGFP-claudin-2 strands (box 1) are replaced by amorphous aggregates at sites of asymmetric mCherry-claudin-4 expression (box 2), suggesting that cis interactions are sufficient for claudin-4-mediated disruption of claudin-2 strands. Maximum projection images representative of 8 independent experiments. Scale: 1 µm. Data are presented as mean ± SD and included in the Source Data file.