---
title: "Lysosomes drive macrophage dissolution of cholesterol crystals revealed by multimodal 3D imaging"
id: "biorxiv-21-multimodal-3d-imaging-reveals-a-central-role-for-lysosomes-indissolution-of"
canonical_url: "https://medichelpline.com/clinical-feed/biorxiv-21-multimodal-3d-imaging-reveals-a-central-role-for-lysosomes-indissolution-of"
content_type: "clinical_feed_article"
specialty: "Cardiology"
source_name: "bioRxiv (Biomedical Preprints)"
source_url: "https://www.biorxiv.org/content/10.64898/2026.09.22.753145v1?rss=1"
published_at: "2026-09-23T09:50:14.000Z"
evidence_level: "Verified Feed"
license: "CC-BY-NC-4.0 / Informational Use"
---
# Lysosomes drive macrophage dissolution of cholesterol crystals revealed by multimodal 3D imaging
## Provenance & Clinical Metadata
- **Canonical URL:** https://medichelpline.com/clinical-feed/biorxiv-21-multimodal-3d-imaging-reveals-a-central-role-for-lysosomes-indissolution-of
- **Specialty:** [Cardiology](https://medichelpline.com/clinical-feed/cardiology.md)
- **Primary Source:** bioRxiv (Biomedical Preprints)
- **Source URL:** [Original Journal Publication](https://www.biorxiv.org/content/10.64898/2026.09.22.753145v1?rss=1)
- **Published At:** 2026-09-23T09:50:14.000Z
- **Evidence Rating:** Verified Feed
## Executive GIST (TL;DR)
- The study used multimodal imaging to examine how macrophages process **cholesterol crystals (CCs)**, focusing on intracellular pathways responsible for CC dissolution. - Researchers incorporated the fluorescent cholesterol analogue **TopFluor-Cholesterol (TF-Chol)** into CCs to visualize uptake and trafficking in macrophages by correlative fluorescence and soft X-ray microscopy. - Quantitative **3D live-cell imaging** showed that CCs are processed in **late endosomes and lysosomes (LE/Lys)** and that this processing leads to formation of TF-Chol–containing **lipid droplets (LDs)** over time. - Pharmacologic perturbations altered CC handling: inhibition of lysosomal sterol export with **U18666A** led to TF-Chol accumulation in LE/Lys, while inhibition of lysosomal acidification with **bafilomycin A1** reduced CC dissolution. - Large CCs contacting macrophages were processed via **lysosomal exocytosis**, followed by both extracellular and intracellular degradation; a novel assay plus 3D image processing supported these observations. - Treatment with a fluorescent **cyclodextrin (CD)** accelerated CC dissolution and increased TF-Chol–enriched LD formation; most fluorescent CD co-localized with LE/Lys markers, suggesting intracellular delivery to LE/Lys contributes to crystal dissolution. - The intrinsically fluorescent sterol **dehydroergosterol (DHE)**, which can form aggregates and crystals, was used with fluorescence spectroscopy and specialized ultraviolet microscopy to show that CD enhances dissolution of DHE crystals in vitro and in cells. - Overall, the data identify the **lysosomal pathway** as central to CC dissolution, cholesterol trafficking, and efflux in macrophages. - The article is a preprint and has not been peer reviewed. Funding sources and competing interest statements are reported in the source; the authors declared no competing interests.
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Egebjerg, Christoph Pratsch, Stephan Werner, Gerd Schneider, Tido Willms, Peter Müller, Sergey Kapishnikov, [ View ORCID Profile](http://orcid.org/0000-0003-4995-9709)Daniel Wüstner doi: https://doi.org/10.64898/2026.09.22.753145 This article is a preprint and has not been certified by peer review [[what does this mean?](https://www.biorxiv.org/about/FAQ#unrefereed)]. Vibeke Akkerman 1 Department of Biochemistry and Molecular Biology, University of Southern Denmark; * [Find this author on Google Scholar](https://www.biorxiv.org/lookup/google-scholar?link_type=googlescholar&gs_type=author&author%5B0%5D=Vibeke%2BAkkerman%2B "Open in new tab") * [Find this author on PubMed](https://www.biorxiv.org/lookup/external-ref?access_num=Akkerman%20V&link_type=AUTHORSEARCH "Open in new tab") * [Search for this author on this site](https://www.biorxiv.org/search/author1%3AVibeke%2BAkkerman%2B) * [ORCID record for Vibeke Akkerman](http://orcid.org/0000-0002-5103-1240 "Open in new tab") Alice Dupont Juhl 1 Department of Biochemistry and Molecular Biology, University of Southern Denmark; * [Find this author on Google Scholar](https://www.biorxiv.org/lookup/google-scholar?link_type=googlescholar&gs_type=author&author%5B0%5D=Alice%2BDupont%2BJuhl%2B "Open in new tab") * [Find this author on PubMed](https://www.biorxiv.org/lookup/external-ref?access_num=Juhl%20AD&link_type=AUTHORSEARCH "Open in new tab") * [Search for this author on this site](https://www.biorxiv.org/search/author1%3AAlice%2BDupont%2BJuhl%2B) Jacob M. Egebjerg 1 Department of Biochemistry and Molecular Biology, University of Southern Denmark; * [Find this author on Google Scholar](https://www.biorxiv.org/lookup/google-scholar?link_type=googlescholar&gs_type=author&author%5B0%5D=Jacob%2BM.%2BEgebjerg%2B "Open in new tab") * [Find this author on PubMed](https://www.biorxiv.org/lookup/external-ref?access_num=Egebjerg%20JM&link_type=AUTHORSEARCH "Open in new tab") * [Search for this author on this site](https://www.biorxiv.org/search/author1%3AJacob%2BM.%2BEgebjerg%2B) * [ORCID record for Jacob M. Egebjerg](http://orcid.org/0009-0001-1451-9116 "Open in new tab") Christoph Pratsch 2 Department of X-Ray Microscopy, Helmholtz-Zentrum Berlin; * [Find this author on Google Scholar](https://www.biorxiv.org/lookup/google-scholar?link_type=googlescholar&gs_type=author&author%5B0%5D=Christoph%2BPratsch%2B "Open in new tab") * [Find this author on PubMed](https://www.biorxiv.org/lookup/external-ref?access_num=Pratsch%20C&link_type=AUTHORSEARCH "Open in new tab") * [Search for this author on this site](https://www.biorxiv.org/search/author1%3AChristoph%2BPratsch%2B) Stephan Werner 2 Department of X-Ray Microscopy, Helmholtz-Zentrum Berlin; * [Find this author on Google Scholar](https://www.biorxiv.org/lookup/google-scholar?link_type=googlescholar&gs_type=author&author%5B0%5D=Stephan%2BWerner%2B "Open in new tab") * [Find this author on PubMed](https://www.biorxiv.org/lookup/external-ref?access_num=Werner%20S&link_type=AUTHORSEARCH "Open in new tab") * [Search for this author on this site](https://www.biorxiv.org/search/author1%3AStephan%2BWerner%2B) Gerd Schneider 2 Department of X-Ray Microscopy, Helmholtz-Zentrum Berlin; * [Find this author on Google Scholar](https://www.biorxiv.org/lookup/google-scholar?link_type=googlescholar&gs_type=author&author%5B0%5D=Gerd%2BSchneider%2B "Open in new tab") * [Find this author on PubMed](https://www.biorxiv.org/lookup/external-ref?access_num=Schneider%20G&link_type=AUTHORSEARCH "Open in new tab") * [Search for this author on this site](https://www.biorxiv.org/search/author1%3AGerd%2BSchneider%2B) Tido Willms 3 Department of Biology, Humboldt University Berlin; * [Find this author on Google Scholar](https://www.biorxiv.org/lookup/google-scholar?link_type=googlescholar&gs_type=author&author%5B0%5D=Tido%2BWillms%2B "Open in new tab") * [Find this author on PubMed](https://www.biorxiv.org/lookup/external-ref?access_num=Willms%20T&link_type=AUTHORSEARCH "Open in new tab") * [Search for this author on this site](https://www.biorxiv.org/search/author1%3ATido%2BWillms%2B) Peter Müller 3 Department of Biology, Humboldt University Berlin; * [Find this author on Google Scholar](https://www.biorxiv.org/lookup/google-scholar?link_type=googlescholar&gs_type=author&author%5B0%5D=Peter%2BM%C3%BCller%2B "Open in new tab") * [Find this author on PubMed](https://www.biorxiv.org/lookup/external-ref?access_num=M%C3%BCller%20P&link_type=AUTHORSEARCH "Open in new tab") * [Search for this author on this site](https://www.biorxiv.org/search/author1%3APeter%2BM%25C3%25BCller%2B) Sergey Kapishnikov 4 SiriusXT Limited, Dublin * [Find this author on Google Scholar](https://www.biorxiv.org/lookup/google-scholar?link_type=googlescholar&gs_type=author&author%5B0%5D=Sergey%2BKapishnikov%2B "Open in new tab") * [Find this author on PubMed](https://www.biorxiv.org/lookup/external-ref?access_num=Kapishnikov%20S&link_type=AUTHORSEARCH "Open in new tab") * [Search for this author on this site](https://www.biorxiv.org/search/author1%3ASergey%2BKapishnikov%2B) Daniel Wüstner 1 Department of Biochemistry and Molecular Biology, University of Southern Denmark; * [Find this author on Google Scholar](https://www.biorxiv.org/lookup/google-scholar?link_type=googlescholar&gs_type=author&author%5B0%5D=Daniel%2BW%C3%BCstner%2B "Open in new tab") * [Find this author on PubMed](https://www.biorxiv.org/lookup/external-ref?access_num=W%C3%BCstner%20D&link_type=AUTHORSEARCH "Open in new tab") * [Search for this author on this site](https://www.biorxiv.org/search/author1%3ADaniel%2BW%25C3%25BCstner%2B) * [ORCID record for Daniel Wüstner](http://orcid.org/0000-0003-4995-9709 "Open in new tab") * For correspondence: wuestner@bmb.sdu.dk * [Abstract](https://www.biorxiv.org/content/10.64898/2026.09.22.753145v1)[](https://www.biorxiv.org/panels_ajax_tab/biorxiv_tab_art/node:5802714/1) * [Info/History](https://www.biorxiv.org/content/10.64898/2026.09.22.753145v1.article-info)[](https://www.biorxiv.org/panels_ajax_tab/biorxiv_tab_info/node:5802714/1) * [Metrics](https://www.biorxiv.org/content/10.64898/2026.09.22.753145v1.article-metrics)[](https://www.biorxiv.org/panels_ajax_tab/article_tab_metrics/node:5802714/1) * [Supplementary material](https://www.biorxiv.org/content/10.64898/2026.09.22.753145v1.supplementary-material)[](https://www.biorxiv.org/panels_ajax_tab/biorxiv_tab_data/node:5802714/1) * [ Preview PDF](https://www.biorxiv.org/content/10.64898/2026.09.22.753145v1.full.pdf+html)[](https://www.biorxiv.org/panels_ajax_tab/biorxiv_tab_pdf/node:5802714/1) ![Loading](https://www.biorxiv.org/sites/all/modules/contrib/panels_ajax_tab/images/loading.gif) ## Abstract Formation of cholesterol crystals (CCs) is a key event during the development of atherosclerosis, but the molecular mechanisms of their degradation within cells are poorly understood. By incorporating the fluorescent cholesterol analogue TopFluor-Cholesterol (TF-Chol) into CCs, we were able to visualize the uptake of CCs in macrophages using correlative fluorescence and soft X-ray microscopy. Using quantitative 3D live-cell imaging, we show that CCs are processed in late endosomes and lysosomes (LE/Lys), resulting in formation of TF-Chol containing lipid droplets (LDs) over time. Inhibition of lysosomal sterol export with U18666A caused accumulation of TF-Chol in LE/Lys, and inhibition of lysosomal acidification with bafilomycin A1 led to reduced dissolution of the CCs. Using a novel assay combined with 3D image processing, we show that large CCs in contact with macrophages are processed via lysosomal exocytosis followed by extracellular and intracellular degradation of CCs. Treating macrophages with a fluorescent version of cyclodextrin (CD) promoted the dissolution of CCs and enhanced the formation of LDs enriched with TF-Chol. The majority of fluorescent CD co-localized with a marker for LE/Lys during this process, suggesting that intracellular delivery to LE/Lys may contribute to the dissolution of CCs. Dehydroergosterol (DHE) is an intrinsically fluorescent sterol closely mimicking the properties and behavior of cholesterol. DHE is known to self-associate into aggregates and crystals, and by using fluorescence spectroscopy and specialized ultraviolet (UV) microscopy, we found that CD enhances the dissolution of DHE crystals in vitro and in cells. Together, our findings highlight the lysosomal pathway as responsible for dissolution of CCs, cholesterol trafficking, and efflux in macrophages. ### Competing Interest Statement The authors have declared no competing interest. ## Funder Information Declared Danish National Research Foundation, https://ror.org/00znyv691, 2032-00136B Carlsberg Foundation, https://ror.org/01kpjmx04, CF24-1904 Novo Nordisk Foundation, https://ror.org/04txyc737, NNF18SA0032928 European Union's Horizon 2020 researchand innovation programme, 101017116 Copyright The copyright holder for this preprint is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made available under a [CC-BY-NC-ND 4.0 International license](http://creativecommons.org/licenses/by-nc-nd/4.0/). bioRxiv and medRxiv thank the following for their generous financial support: > The Chan Zuckerberg Initiative, Cold Spring Harbor Laboratory, the Sergey Brin Family Foundation, California Institute of Technology, Centre National de la Recherche Scientifique, Fred Hutchinson Cancer Center, Imperial College London, Massachusetts Institute of Technology, Stanford University, The University of Edinburgh, University of Washington, and Vrije Universiteit Amsterdam. [Donate to openRxiv ](https://www.zeffy.com/en-US/donation-form/donate-to-make-a-difference-10981) [ Back to top](https://www.biorxiv.org/content/10.64898/2026.09.22.753145v1?rss=1#page) [ Previous](https://www.biorxiv.org/content/10.64898/2026.09.21.753355v1 "Loss of TCA cycle turning promotes stem cell function") Posted September 23, 2026. [ Download PDF](https://www.biorxiv.org/content/10.64898/2026.09.22.753145v1.full.pdf) Print/Save Options [Download PDF](https://www.biorxiv.org/content/biorxiv/early/2026/09/23/2026.09.22.753145.full.pdf)Full Text & In-line FiguresXML [More Info](https://www.biorxiv.org/about/FAQ#PrintOptions "More Information on Print/Save Options") [Supplementary Material ](https://www.biorxiv.org/content/10.64898/2026.09.22.753145v1.supplementary-material) [ Email](https://www.biorxiv.org/ "Email this Article") [ Share](https://www.biorxiv.org/) Multimodal 3D imaging reveals a central role for lysosomes in dissolution of cholesterol crystals by macrophages Vibeke Akkerman, Alice Dupont Juhl, Jacob M. Egebjerg, Christoph Pratsch, Stephan Werner, Gerd Schneider, Tido Willms, Peter Müller, Sergey Kapishnikov, Daniel Wüstner bioRxiv 2026.09.22.753145; doi: https://doi.org/10.64898/2026.09.22.753145 This article is a preprint and has not been certified by peer review [[what does this mean?](https://www.biorx
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