Accumulation of endogenous free radicals is required to induce titan-like cell formation in Cryptococcus neoformans
Open accessAbstract
Cryptococcus neoformans is an encapsulated yeast able to cause disease (mainly meningoencephalitis) among immunosuppressed patients, mostly HIV+. This yeast can form the so-called titan cells in vivo , which are cells of an abnormally larger size due to an increase in both the capsule and the cell body size (total size reaching between 50–70 μm). This phenomenon can be partially reproduced in vitro to obtain cells of an intermediate size (25–30 μm), which have been denominated titan-like cells. In this work, we have screened 1,520 compounds from the Prestwick Chemical Library and identified off-patent drugs that inhibited titan-like cell formation in vitro . We developed an automated fluorescence-based microscopy assay and identified 64 compounds as possible inhibitors of titan-like cells. We chose 10 of these compounds to perform dose-response curves and confirmed them as inhibitors of titan-like cell formation. Several of the compounds identified had antioxidant properties (i.e., retinoic acid), indicating a possible role of free radicals during titan cell formation. Using fluorescent probes, we found that there was an endogenous accumulation of reactive oxygen species (ROS) during cell growth, which was inhibited in the presence of retinoic acid. Furthermore, we found that during titanization, there were significant changes in the mitochondria, which is the main organelle where ROS are produced. We hypothesize that an intracellular increase of free radicals at the mitochondria might be a triggering signal to induce titanization. IMPORTANCE: Cryptococcus neoformans is an excellent model to investigate fungal pathogenesis. This yeast can produce “titan cells,” which are cells of an abnormally larger size that contribute to the persistence of the yeast in the host. In this work, we have used a new approach to characterize them by identifying drugs that inhibit this process. We have used a repurposing off-patent drug library, combined with an automatic method to image and analyze fungal cell size. In this way, we have identified many compounds that inhibit this transition. Interestingly, several compounds were antioxidants, allowing us to confirm that endogenous ROS and mitochondrial changes are important for titan cell formation. This work provides new evidence of the mechanisms required for titanization. Furthermore, the future characterization of the inhibitory mechanisms of the identified compounds by the scientific community will contribute to better understand the role of titan cells in virulence.
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The process of titan cell production of aneuploid daughters was associated with intracellular ROS accumulation and mitochondrial responses.
“This process was associated with intracellular ROS accumulation and mitochondrial responses [141].”
From Insight into the Mechanisms and Clinical Relevance of Antifungal Heteroresistance (2025) · cites this paper as [141]
The Results section on ROS detection reports intracellular ROS accumulation during titan-like cell formation, and the Discussion reports associated changes in mitochondrial activity. The paper studies titan-like cell formation, however, rather than showing that ROS accumulation or mitochondrial responses are associated specifically with the production of aneuploid daughter cells. Our copy provides no page numbers for these passages. (LLM, conf=0.84)
Experimental — accuracy not yet measuredOne mechanism of C. neoformans heteroresistance is associated with multiple types of aneuploid daughter cells produced by titan cells.
“One of its mechanisms is associated with the multiple types of aneuploid daughter cells produced by titan cells [56,140,141].”
From Insight into the Mechanisms and Clinical Relevance of Antifungal Heteroresistance (2025) · cites this paper as [141]
The Introduction describes titan cells as polyploid and discusses their formation, but does not say they produce aneuploid daughter cells. Searches for aneuploid daughter cells and heteroresistance found no passage connecting titan cells to the claimed resistance mechanism. The indexed copy provides no page numbers for these passages. (LLM, conf=0.75)
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- Authors
- García-Barbazán, I., Torres-Cano, A., García-Rodas, R., Sachse, M., Luque, D., Megías, D., & Zaragoza, O.
- Affiliations
- Instituto de Salud Carlos III (ES)
- Mycology Reference Laboratory, National Centre for Microbiology, Instituto de Salud Carlos III, Majadahonda, Madrid, Spain
- Advanced Optical Microscopy Unit, Central Core Facilities, Instituto de Salud Carlos III, Madrid, Spain
- Venue
- mBio
- Publisher
- American Society for Microbiology
- Published
- 2024-01-16
- Volume
- 15
- Issue
- 1
- Pages
- e0254923-e0254923
- ISSN
- 2150-7511 (ISSN-L)
- PMC
- PMC10790760
- OpenAlex
- W4389558743
- Cited by
- 14
- References
- 73
- Citations by year
- 2026: 9 · 2025: 4 · 2024: 1
- FWCI
- 2.78
- Type
- journal article
- Language
- en
- Open access
- gold
- doi.org
- Licence
- Locations
- mBio (publishedVersion, cc-by)
- PubMed Central (publishedVersion, cc-by)
- PubMed (publishedVersion)
- UNSWorks (University of New South Wales, Sydney, Australia) (submittedVersion, cc-by-nc-nd)
- DOAJ (DOAJ: Directory of Open Access Journals) (submittedVersion, cc-by-sa)
- Funders
- MEC | Agencia Estatal de Investigación (PID2020-114546RB by by MCIN/AEI/ 10.13039/501100011033)
- Ministerio de Ciencia e Innovación (SAF2017-86912-R, FPI-PRE2018-083436, FPI-PRE2021-099486, IJCI-2015-25683)
- Topics
- Health Sciences › Medicine › Epidemiology › Fungal Infections and Studies
- Health Sciences › Medicine › Infectious Diseases › Antifungal resistance and susceptibility
- Life Sciences › Biochemistry, Genetics and Molecular Biology › Molecular Biology › Fungal and yeast genetics research
- Keywords
- Cryptococcus neoformans, titan cells, reactive oxygen species, free radicals, mitochondrial dysfunction, retinoic acid, antioxidants, drug repurposing, fungal virulence, meningoencephalitis, HIV-associated cryptococcosis
- MeSH terms
- Cryptococcosis, Cryptococcus neoformans, Saccharomyces cerevisiae, Virulence
About these checks
Registry, retraction, Cites and statistics results are computed, not judged, with one exception: a printed reference that no index lists is read once by a model before it counts as a citation error. Claims are extracted by a model and kept only when their quote appears verbatim in the text; each cited source is then checked, with a model’s help, for whether it says what the claim relies on. Those verdicts can be wrong — read the claim’s quote and the cited source before relying on one.
Statistics and arithmetic are recomputed, not judged.
Confidence bands are shown only for fields calibrated on our annotated test set. None is calibrated yet, so no band is shown.
What counts as a claim ›How we measure confidence ›How we check this ›
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