Cited by 160Antifungal resistance and susceptibilityInfection Control in HealthcareBacterial biofilms and quorum sensing

Candida auris Forms High-Burden Biofilms in Skin Niche Conditions and on Porcine Skin

Open access
Horton, Mark V., Johnson, Chad J., Kernien, John F., Patel, Tarika D., Lam, Brandon C., Cheong, J. Z. Alex, Meudt, Jennifer J., Shanmuganayagam, Dhanansayan +2 more·2020·mSphere

Abstract

The emerging fungal pathogen Candida auris causes invasive infections and is spreading in hospitals worldwide. Why this species exhibits the capacity to transfer efficiently among patients is unknown. Our findings reveal that C. auris forms high-burden biofilms in conditions mimicking sweat on the skin surface. These adherent biofilm communities persist in environmental conditions expected in the hospital setting. Using a pig skin model, we show that C. auris also forms high-burden biofilm structures on the skin surface. Identification of this mode of growth sheds light on how this recently described pathogen persists in hospital settings and spreads among patients.

AI snapshot

Written from the paper’s full text.

TL;DR

Candida auris forms high-burden biofilms in synthetic human sweat and on ex vivo porcine skin, exceeding Candida albicans under these skin-niche conditions. These biofilms persist through sweat evaporation and desiccation, suggesting a mechanism for hospital persistence and spread.

Key findings

  • In synthetic human axillary sweat, C. auris produced biofilms with 10-fold greater burden than C. albicans after 24 h (P < 0.05), while in RPMI-MOPS C. albicans biofilm biomass was 3-fold greater than C. auris (P < 0.05).
  • In concentrated synthetic sweat, C. auris biofilms were 10-fold greater than C. albicans at 1.5× and 4-fold greater at 2.5×. After 24 h desiccation, C. auris burden was approximately 30-fold greater than C. albicans at 7 days; at 14 days C. auris grew readily while C. albicans biofilms were not viable.
  • On porcine skin, C. auris reached a burden nearly 15-fold greater than C. albicans and replicated nearly 100-fold above the initial inoculum (P < 0.05); scanning electron microscopy showed multilayer yeast-cell biofilms without tissue invasion.

Methods

  • Biofilm assays used C. albicans SC5314 and C. auris strains from the CDC, including B11203, grown in RPMI-MOPS or synthetic human axillary sweat medium; burden was measured by OD600, and architecture was imaged by scanning electron microscopy.
  • Desiccation assays grew biofilms for 24 h, desiccated them in a laminar flow hood for 24 h, then assessed viable burden after 7 or 14 days by plating on YPD agar.
  • The ex vivo skin model used full-thickness 12-mm skin samples from Wisconsin Miniature Swine, placed in DMEM with 10% FBS and a paraffin barrier, inoculated on the epidermal surface with Candida in synthetic sweat medium, incubated 24 h, and assessed by CFU counts and scanning electron microscopy.

Limitations

  • The paper states no limitations.
  • Evident from the text: the main skin model is ex vivo porcine skin, not human skin, and the in vitro work uses synthetic sweat medium rather than native human sweat.
  • Evident from the text: most experiments use C. auris strain B11203, although a CDC strain set spanning clades was tested for initial biofilm formation; mechanisms of desiccation resistance and biofilm formation are not determined.

Model-written summary. Check it against the paper before relying on it.

Integrity signals

Registry

Crossref · 10.1128/msphere.00910-19

Cites

33 of 33 references verified0 citation errors · 0 not checkable

Statistics

Nothing to recompute.

Claims the paper makes

8 of 20 cited sources checked · 6 supported · 0 flagged · accuracy not yet measured

  • Findingown data

    On porcine skin, C. auris forms multilayer biofilms composed of yeast cells.

    “Higher magnification imaging showed the formation of multilayer biofilms composed of yeast cells.”
    Quote in the paperSubstantial claim
  • Findingown data

    In synthetic sweat medium mimicking human axillary sweat, after 24 h, C. auris produced biofilms with 10-fold greater burden than C. albicans.

    “After 24 h of incubation, C. auris produced dense biofilms with 10-fold greater burden than the biofilms formed by C. albicans (Fig. 1a) (P < 0.05).”
    Quote in the paperSubstantial claim
  • Findingown data

    In RPMI-MOPS, the biomass of biofilms formed by C. albicans was 3-fold greater than for C. auris.

    “For example, in RPMI-MOPS (morpholinepropanesulfonic acid), the biomass of biofilms formed by C. albicans was 3-fold greater than for C. auris (Fig. 1a) (P < 0.05), consistent with prior studies demonstrating lower biofilm burdens for this species (8, 9).”
    Quote in the paperSubstantial claim
  • Findingown data

    Even in 2.5×-concentrated synthetic sweat, C. auris exhibited the capacity to proliferate as a biofilm.

    “Even in 2.5×-concentrated sweat, C. auris exhibited the capacity to proliferate as a biofilm.”
    Quote in the paperSubstantial claim
  • Findingown data

    C. auris forms high-burden biofilms in conditions mimicking sweat on the skin surface.

    “Our findings reveal that C. auris forms high-burden biofilms in conditions mimicking sweat on the skin surface.”
    Quote in the paperSubstantial claim
  • May combine two claims. 12
  • Classified as probably not a substantial claim. 6
  • May combine two claims.

    Interpretationown data

    These observations help explain C. auris's propensity for colonizing skin, persisting on hospital surfaces, and spreading efficiently in the hospital setting.

    “These observations help to explain this pathogen’s propensity for colonizing skin, persisting on hospital surfaces, and spreading efficiently in the hospital setting (4, 19).”
    Quote in the paperSubstantial claim2 of 2 sources checked
    • cites [4] Schelenz S, Hagen F, Rhodes JL, Abdolrasouli A, Chowdhary A, Hall A, Ryan L, Shackleton J, Trimlett R, Meis JF, Armstrong-James D, Fisher MC 2016 First hospital outbreak of the globally emerging Candida auris in a European hospital. Antimicrob Resist Infect Control 5: 35. doi: 10.1186/s13756-016-0132-5. 27777756 PMC5069812 source
    • cites [19] Vallabhaneni S, Kallen A, Tsay S, Chow N, Welsh R, Kerins J, Kemble SK, Pacilli M, Black SR, Landon E, Ridgway J, Palmore TN, Zelzany A, Adams EH, Quinn M, Chaturvedi S, Greenko J, Fernandez R, Southwick K, Furuya EY, Calfee DP, Hamula C, Patel G, Barrett P, MSD, Lafaro P, Berkow EL, Moulton-Meissner H, Noble-Wang J, Fagan RP, Jackson BR, Lockhart SR, Litvintseva AP, Chiller TM 2016 Investigation of the first seven reported cases of Candida auris, a globally emerging invasive, multidrug-resistant fungus - United States, May 2013-August 2016. MMWR Morb Mortal Wkly Rep 65: 1234– 1237. doi: 10.15 source

      The main report (page unavailable in our copy) documents skin colonization weeks to months after infection and C. auris on multiple surfaces in one hospital room. It says these findings suggest spread in health care settings is possible, but transmission was not definitively documented; it does not establish how long the organism persisted on surfaces or that it spread efficiently. (LLM, conf=0.86)

      Experimental — accuracy not yet measured
  • Classified as probably not a substantial claim. May combine two claims.

    Premisecitation

    Circulating C. auris strains vary genetically and cluster into at least 4 distinct clades.

    “Circulating C. auris strains vary genetically and cluster into at least 4 distinct clades (1).”
    Quote in the paperSubstantial claim1 of 1 sources checked
    • cites [1] Lockhart SR, Etienne KA, Vallabhaneni S, Farooqi J, Chowdhary A, Govender NP, Colombo AL, Calvo B, Cuomo CA, Desjardins CA, Berkow EL, Castanheira M, Magobo RE, Jabeen K, Asghar RJ, Meis JF, Jackson B, Chiller T, Litvintseva AP 2017 Simultaneous emergence of multidrug-resistant Candida auris on 3 continents confirmed by whole-genome sequencing and epidemiological analyses. Clin Infect Dis 64: 134– 140. doi: 10.1093/cid/ciw691. 27988485 PMC5215215 source
  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Premisecitation

    Day and colleagues showed that C. auris exhibited a unique stress resistance profile compared to other Candida spp.

    “However, studies by Day and colleagues characterized the responses of Candida to a variety of stresses, including high salinity, and showed that C. auris exhibited a unique stress resistance profile compared to those of other Candida spp. (22).”
    Quote in the paperNot a substantial claim1 of 1 sources checked
    • cites [22] Day AM, McNiff MM, da Silva Dantas A, Gow NAR, Quinn J 2018 Hog1 regulates stress tolerance and virulence in the emerging fungal pathogen Candida auris. mSphere 3: e00506-18. doi: 10.1128/mSphere.00506-18. 30355673 PMC6200985 source
  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Premisecitation

    The stress responses studied by Day and colleagues involved the evolutionarily conserved Hog1 stress-activated protein kinase (SAPK).

    “Despite the different profiles, these responses involved the evolutionarily conserved Hog1 stress-activated protein kinase (SAPK).”
    Quote in the paperNot a substantial claim1 of 1 sources checked
    • cites [22] Day AM, McNiff MM, da Silva Dantas A, Gow NAR, Quinn J 2018 Hog1 regulates stress tolerance and virulence in the emerging fungal pathogen Candida auris. mSphere 3: e00506-18. doi: 10.1128/mSphere.00506-18. 30355673 PMC6200985 source
  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Premisecitation

    A prior study has shown C. auris to survive dry conditions for up to 2 weeks.

    “as a prior study has shown this species to survive dry conditions for up to 2 weeks (21).”
    Quote in the paperNot a substantial claim1 of 1 sources checked
    • cites [21] Welsh RM, Bentz ML, Shams A, Houston H, Lyons A, Rose LJ, Litvintseva AP 2017 Survival, persistence, and isolation of the emerging multidrug-resistant pathogenic yeast Candida auris on a plastic health care surface. J Clin Microbiol 55: 2996– 3005. doi: 10.1128/JCM.00921-17. 28747370 PMC5625385 source
  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Premisecitation

    The karyotypic plasticity displayed by C. auris in response to stressors may contribute to this organism's ability to adapt and persist in the skin niche.

    “In addition, the karyotypic plasticity displayed by C. auris in response to stressors may contribute to this organism’s ability to adapt and persist in the skin niche (23).”
    Quote in the paperNot a substantial claim1 of 1 sources checked
    • cites [23] Bravo Ruiz G, Ross ZK, Holmes E, Schelenz S, Gow NAR, Lorenz A 2019 Rapid and extensive karyotype diversification in haploid clinical Candida auris isolates. Curr Genet 65: 1217– 1228. doi: 10.1007/s00294-019-00976-w. 31020384 PMC6744574 source

      The Results section (‘C. auris undergoes karyotype rearrangements in stress conditions’; page unavailable) documents stress-induced karyotype changes, and the Discussion suggests such changes might aid fitness adaptation. The Introduction describes C. auris as a skin colonizer, but the paper does not link karyotypic plasticity to adaptation or persistence in the skin niche. (LLM, conf=0.88)

      Experimental — accuracy not yet measured
  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Premisecitation

    Individual C. auris strains appear to have arisen independently in remote geographic locations.

    “What is striking about C. auris is that individual strains appear to have arisen independently in remote geographic locations (1).”
    Quote in the paperNot a substantial claim1 of 1 sources checked
    • cites [1] Lockhart SR, Etienne KA, Vallabhaneni S, Farooqi J, Chowdhary A, Govender NP, Colombo AL, Calvo B, Cuomo CA, Desjardins CA, Berkow EL, Castanheira M, Magobo RE, Jabeen K, Asghar RJ, Meis JF, Jackson B, Chiller T, Litvintseva AP 2017 Simultaneous emergence of multidrug-resistant Candida auris on 3 continents confirmed by whole-genome sequencing and epidemiological analyses. Clin Infect Dis 64: 134– 140. doi: 10.1093/cid/ciw691. 27988485 PMC5215215 source
  • May combine two claims.

    Findingown data

    In RPMI-MOPS, C. albicans forms dense biofilms composed primarily of hyphae, while C. auris forms a monolayer of yeast.

    “In this tissue culture medium, C. albicans formed dense biofilms composed primarily of hyphae, while biofilms formed by C. auris appeared as a monolayer of yeast.”
    Quote in the paperSubstantial claim
  • May combine two claims.

    Findingown data

    Biofilm growth by the majority of C. auris strains paralleled strain B11203, with burdens reaching approximately 10-fold that of C. albicans.

    “Biofilm growth by the majority of the strains paralleled that observed for C. auris B11203 (listed as strain 5), with burdens reaching approximately 10-fold that of C. albicans (see Fig. S1).”
    Quote in the paperSubstantial claim
  • May combine two claims.

    Findingown data

    On porcine skin, C. auris exhibited enhanced growth compared to C. albicans, propagating to a burden nearly 15-fold greater than C. albicans and replicating nearly 100-fold above the initial inoculum.

    “Consistent with our in vitro observations, C. auris exhibited enhanced growth compared to that of C. albicans in this environment, with C. auris propagating to a burden nearly 15-fold greater than C. albicans and replicating nearly 100-fold above the initial inoculum (Fig. 2b) (P < 0.05).”
    Quote in the paperSubstantial claim
  • May combine two claims.

    Findingown data

    On porcine skin, C. albicans showed little fungal growth and multilayer aggregates were not observed.

    “In contrast, imaging of porcine skins inoculated with C. albicans showed little fungal growth. Multilayer aggregates were not observed.”
    Quote in the paperSubstantial claim
  • May combine two claims.

    Findingown data

    At 14 days postdesiccation, C. auris grew readily from biofilms, while C. albicans biofilms were not viable.

    “At 14 days postdesiccation, C. auris grew readily from biofilms, while C. albicans biofilms were not viable (Fig. 1e).”
    Quote in the paperSubstantial claim
  • May combine two claims.

    Findingown data

    In 2.5×-concentrated synthetic sweat, C. auris biofilm burdens were below those formed in less-concentrated sweat but were still 4-fold greater than C. albicans biofilms.

    “The burdens of these biofilms were below those formed in the less-concentrated sweat but were still 4-fold greater than the biofilms formed by C. albicans.”
    Quote in the paperSubstantial claim
  • May combine two claims.

    Premisecitation

    Pig skin shares many characteristics with human skin, including similarities in the thickness of skin layers, the types and distribution of skin cells, and the mechanisms of skin repair.

    “Pig skin was selected based on the many characteristics it shares with human skin, including similarities in the thickness of skin layers, the types and distribution of skin cells, and the mechanisms of skin repair (15–18).”
    Quote in the paperSubstantial claim0 of 4 sources checked
    • cites [15] Liu Y, Chen JY, Shang HT, Liu CE, Wang Y, Niu R, Wu J, Wei H 2010 Light microscopic, electron microscopic, and immunohistochemical comparison of Bama minipig (Sus scrofa domestica) and human skin. Comp Med 60: 142– 148. 20412690 PMC2855042
    • cites [16] Eaglstein WH, Mertz PM 1978 New methods for assessing epidermal wound healing: the effects of triamcinolone acetonide and polyethelene film occlusion. J Invest Dermatol 71: 382– 384. doi: 10.1111/1523-1747.ep12556814. 722115 source
    • cites [17] Sullivan TP, Eaglstein WH, Davis SC, Mertz P 2001 The pig as a model for human wound healing. Wound Repair Regen 9: 66– 76. doi: 10.1046/j.1524-475x.2001.00066.x. 11350644 source
    • cites [18] Summerfield A, Meurens F, Ricklin ME 2015 The immunology of the porcine skin and its value as a model for human skin. Mol Immunol 66: 14– 21. doi: 10.1016/j.molimm.2014.10.023. 25466611 source
  • May combine two claims.

    Interpretationown data

    Growth in synthetic sweat medium and on porcine skin allows C. auris to form dense biofilms that flourish in conditions of evaporation and resist desiccation.

    “Our findings demonstrate that growth in synthetic sweat medium and on porcine skin allows C. auris to form dense biofilms that flourish in conditions of evaporation and resist desiccation.”
    Quote in the paperSubstantial claim
  • May combine two claims.

    Interpretationown data

    The lack of tissue invasion supports the use of porcine skin to model human skin colonization, as this process does not involve tissue destruction by fungi.

    “This lack of invasion supports the use of porcine skin to model human skin colonization, as this process does not involve tissue destruction by fungi.”
    Quote in the paperSubstantial claim
    • Rests on an assumption: Human skin colonization by C. auris does not involve tissue destruction by fungi.
  • May combine two claims.

    Interpretationown data

    Growth of C. auris in the ex vivo porcine skin model mimics what is observed clinically, an enhanced capacity for skin colonization.

    “Growth of C. auris in this ex vivo model mimics what is observed clinically, an enhanced capacity for skin colonization.”
    Quote in the paperSubstantial claim
17 of 17
  • The paper relies on this without support.

    Human skin colonization by C. auris does not involve tissue destruction by fungi.

    “This lack of invasion supports the use of porcine skin to model human skin colonization, as this process does not involve tissue destruction by fungi.”
1 of 1

Claims other papers make about this one

Each is checked against this paper’s own text: does it say what the citing paper says it does?

  • Clinical features of C. auris infections include biofilm persistence on abiotic surfaces and skin, immune evasion, and antifungal drug resistance.

    “The clinical features of these infections include biofilm persistence on abiotic surfaces and skin, immune evasion, and antifungal drug resistance (7–12).”

    From Dual function of <i>Candida auris</i> mannosyltransferase, MNT5, in biofilm community protection from antifungal therapy and the host (2025) · cites this paper as [12]

    The Introduction and Discussion (page unavailable) describe C. auris colonizing skin, persisting on hospital surfaces, and forming biofilms under skin-like conditions. The paper does not establish immune evasion or antifungal drug resistance as clinical features; searching those terms yielded references to other studies, not findings reported in the paper’s text. (LLM, conf=0.87)

    Experimental — accuracy not yet measured
  • In an in vitro artificial human axillary sweat medium, C. auris achieves a fungal burden 10 times greater than C. albicans and persists during colonization for over 14 days, whereas C. albicans does not.

    “In an in vitro model using an artificial human axillary sweat medium, C. auris achieves a fungal burden 10 times greater than Candida albicans and persists during colonization for over 14 days, which was not observed in C. albicans18.”

    From Adhesin Als4112 promotes Candida auris skin colonization through interactions with keratinocytes and extracellular matrix proteins (2025) · cites this paper as [18]

    The Results section (“C. auris forms high-burden biofilms in skin milieu”) reports a 10-fold greater C. auris biofilm burden than C. albicans in synthetic sweat medium. The Results section on desiccation reports viable C. auris at 14 days postdesiccation, while C. albicans biofilms were not viable. It does not establish persistence for over 14 days or describe that result as persistence during colonization. (LLM, conf=0.88)

    Experimental — accuracy not yet measured
  • C. auris can grow and persist for extensive periods under skin-like conditions in simulation studies and animal models.

    “Several simulation studies and animal models have demonstrated that C. auris can grow and persist for extensive periods under skin-like conditions.”

    From <i>Candida auris</i> metabolism and growth preferences in physiologically relevant skin-like conditions (2026) · cites this paper as [21]

    The Results report growth on ex vivo porcine skin after 24 hours and survival of biofilms formed in synthetic sweat medium for up to 14 days after desiccation. These findings support growth and persistence under simulated skin-like conditions, but the paper's porcine-skin experiment is not a live-animal model, and the extended persistence was shown in the biofilm experiment rather than on porcine skin. (LLM, conf=0.88)

    Experimental — accuracy not yet measured

Cites

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Authors
Horton, M. V., Johnson, C. J., Kernien, J. F., Patel, T. D., Lam, B. C., Cheong, J. Z. A., Meudt, J. J., Shanmuganayagam, D., Kalan, L. R., & Nett, J. E.
Affiliations
University of Wisconsin–Madison (US)
Venue
mSphere
Published
2020-02-26
Volume
5
Issue
1
ISSN
2379-5042 (ISSN-L)
OpenAlex
W2999046918
Cited by
159
References
33
Citations by year
2026: 34 · 2025: 19 · 2024: 26 · 2023: 31 · 2022: 24 · 2021: 14 · 2020: 11
FWCI
2.41
Type
journal article
Language
en
Open access
Locations
  • mSphere (publishedVersion, cc-by)
  • PubMed (publishedVersion)
  • PubMed Central (submittedVersion, cc-by)
  • DOAJ (DOAJ: Directory of Open Access Journals) (submittedVersion, cc-by-sa)
  • DOAJ (DOAJ: Directory of Open Access Journals) (submittedVersion)
Funders
  • HHS | National Institutes of Health (K08 AI108727)
  • Burroughs Wellcome Fund (1012299)
Topics
  • Health Sciences › Medicine › Infectious Diseases › Antifungal resistance and susceptibility
  • Health Sciences › Medicine › Infectious Diseases › Infection Control in Healthcare
  • Life Sciences › Biochemistry, Genetics and Molecular Biology › Molecular Biology › Bacterial biofilms and quorum sensing
Keywords
Candida auris, biofilm formation, sweat, porcine skin model, hospital-acquired fungal infection, environmental persistence, emerging fungal pathogens
MeSH terms
Animals, Candida, Skin, Sweat, Swine, Biofilms, In Vitro Techniques

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