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. 2020 May 4;11(5):314. doi: 10.1038/s41419-020-2503-3

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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. 1 — a–c Primary cortical neurons were pretreated with internalization inhibitors for 30 min before treating with 0.5 μM BDTOs for 1 h. AD TauO (a), PSP TauO (b), and DLB TauO (c) internalized into cells ~55%, 60%, and 45%, respectively. The percentage of AF568-tagged tau-positive cells and median fluorescence intensity (right panel) from AD TauO (a), PSP TauO (b), and DLB TauO (c) applications were significantly diminished in a concentration-dependent fashion by Heparin (a HSPG antagonist), whereas Dynasore (a clathrin inhibitor) and Nystatin (a caveolae inhibitor) did not show inhibitory effects. A total of 10,000 cells were analyzed for each condition in triplicate. Error bars show SEM. ***p < 0.001, ****p < 0.0001 vs vehicle control (Veh. Con; 0.02% (v/v) DMSO), ^##p < 0.01, ^###p < 0.001, ^####p < 0.0001 vs TauO-treated. All histograms (right panel) represent integrated median fluorescence intensity of BDTOs in cells pretreated with 26–38 μg/ml Dynasore, 1–200 μg/ml Heparin, or 1–40 μg/ml Nystatin. AF568-tagged tau fibrils were used as the negative control in all experiments. See also Supplementary Fig. S3a. d–g Dextran and BDTOs uptake were associated with HSPG. A total of 20 μg/ml of 70,000 MW lysine-flexible Texas Red-conjugated dextran was applied to cells for 1 h. d Cells were fixed and immunostained for HSPG (green) and a mature neuronal marker (βIII-tubulin, blue). Representative orthogonal images indicated AF488-tagged AD TauO (e), PSP TauO (f), and DLB TauO (g) co-localized to dextran, indicating the HSPG-mediated uptake. Scale bar: 10 μm.