The DOI matches its Crossref record; 215 of the 215 references it cites were found in a registry or an index; 1 of the 170 sources its claims cite were checked for support.
Candida albicans Biofilms and Human Disease
Open accessAbstract
In humans, microbial cells (including bacteria, archaea, and fungi) greatly outnumber host cells. Candida albicans is the most prevalent fungal species of the human microbiota; this species asymptomatically colonizes many areas of the body, particularly the gastrointestinal and genitourinary tracts of healthy individuals. Alterations in host immunity, stress, resident microbiota, and other factors can lead to C. albicans overgrowth, causing a wide range of infections, from superficial mucosal to hematogenously disseminated candidiasis. To date, most studies of C. albicans have been carried out in suspension cultures; however, the medical impact of C. albicans (like that of many other microorganisms) depends on its ability to thrive as a biofilm, a closely packed community of cells. Biofilms are notorious for forming on implanted medical devices, including catheters, pacemakers, dentures, and prosthetic joints, which provide a surface and sanctuary for biofilm growth. C. albicans biofilms are intrinsically resistant to conventional antifungal therapeutics, the host immune system, and other environmental perturbations, making biofilm-based infections a significant clinical challenge. Here, we review our current knowledge of biofilms formed by C. albicans and closely related fungal species.
Overview
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Written from the paper’s full text.
TL;DR
This review summarizes how *Candida albicans* forms biofilms on medical devices and mucosal surfaces, and how these communities resist antifungal drugs and host defenses. It highlights a six-regulator transcriptional network, matrix- and persister-mediated drug resistance, and the clinical burden of biofilm infections.
Key findings
- *C. albicans* biofilm development proceeds through four stages: adherence of yeast cells, proliferation into a basal layer, growth of pseudohyphae/hyphae with extracellular matrix production, and dispersal of yeast-form cells. Mature biofilms typically form within 24 hours in vitro.
- Six master transcriptional regulators (Efg1, Tec1, Bcr1, Ndt80, Brg1, Rob1) are each required for biofilm development in vitro and in rat catheter/denture models; together they directly bind and likely regulate approximately 1,000 target genes. The review counts 50 transcriptional regulators and 101 nonregulatory genes with functionally validated biofilm roles.
- Biofilm resistance involves early and sustained efflux pump upregulation, extracellular matrix components including β-1,3-glucan and extracellular DNA, and metabolically dormant persister cells. Matrix composition is reported as ~55% proteins/glycoproteins, 25% carbohydrates, 15% lipids, and 5% nucleic acids, with over 500 proteins identified.
- Clinical impact: NIH estimates biofilms account for over 80% of microbial infections in the United States; CTG clade species account for 15% of hospital-acquired sepsis and are the fourth most frequent cause of bloodstream infections; over 5 million central venous catheters are placed annually in the United States (1992–2001 data), with biofilm infection in over 50%; estimated 100,000 deaths and $6.5 billion in excess expenditure annually in the United States; disseminated candidiasis mortality approaches 40% in some cases.
Methods
- Narrative review of published literature on *C. albicans* and related fungal biofilms; no new experimental data are reported.
- Synthesizes in vitro biofilm models (e.g., silicone squares, flow cells), in vivo/ex vivo animal models (rat/rabbit catheter, denture stomatitis, vaginal mucosal, bioluminescence), and genetic/genomic/proteomic studies (knockout screens, transcriptional profiling, proteomics).
- Main analysis is qualitative synthesis, including tables of 50 transcriptional regulators and 101 nonregulatory genes with validated biofilm roles.
Limitations
- Stated: little is known about persister cell formation and regulation; virtually nothing is known about interactions among *Candida* species in polymicrobial biofilms; the majority of newly identified biofilm-network target genes remain unstudied; the biofilm network outline is “undoubtedly incomplete”; and biofilm formation is often loosely defined in non-*Candida* fungi.
- Evident: as a review, it does not present primary experimental data, and its conclusions depend on the designs and conditions of cited studies; many mechanistic findings come from in vitro models or non-*C. albicans* species.
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Claims the paper makes
85 Not checked yet · 20 Own interpretation · 11 No basis given · 1 Supported · 1 Own result
Every source the paper cites for this claim was checked and supports it.
In the paper
“Together, these six master regulators directly bind to the promoters of and very likely regulate the expression of approximately 1,000 target genes, some of which are additional transcriptional regulators.”
§ Candida albicans Biofilms and Human Disease > GENETICS OF C. ALBICANS BIOFILM FORMATION > Regulatory Control of C. albicans Biofilm Development
What the sources say
- Supportedcites [140] source
In Results, “Developing transcriptional relationships among biofilm regulators,” the authors estimate 1,061 genes with promoter regions bound by at least one of the six regulators and describe them as likely to be controlled. The same section reports that the regulators bind the promoters of other regulators, supporting the final clause. No page numbers are provided in our copy. (LLM, conf=0.93)
hedgedconditional- Asserted: passes — The authors put it forward as true.
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- Supported
In the paper
“Each master regulator also controls the expression of the other master regulators, resulting in a complex, intertwined regulatory network.”
§ Candida albicans Biofilms and Human Disease > GENETICS OF C. ALBICANS BIOFILM FORMATION > Regulatory Control of C. albicans Biofilm Development
What the sources say
- Not checked yetcites [140] source
plaindirect- Asserted: passes — The authors put it forward as true.
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In the paper
“These studies have revealed that many hundreds of mRNAs and proteins are differentially expressed between biofilms and planktonic cells.”
§ Candida albicans Biofilms and Human Disease > GENETICS OF C. ALBICANS BIOFILM FORMATION
What the sources say
plaincorrelational- Asserted: passes — The authors put it forward as true.
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In the paper
“This network consists of six master transcriptional regulators (Efg1, Tec1, Bcr1, Ndt80, Brg1, and Rob1), each of which is required for biofilm development in both in vitro and in vivo rat catheter and rat denture models (140).”
§ Candida albicans Biofilms and Human Disease > GENETICS OF C. ALBICANS BIOFILM FORMATION > Regulatory Control of C. albicans Biofilm Development
What the sources say
- Not checked yetcites [140] source
plaindirect- Asserted: passes — The authors put it forward as true.
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In the paper
“Biofilm architectures in rat and rabbit central venous and indwelling urinary catheter models and in rat denture stomatitis models also include numerous yeast cells in the basal region, and hyphae and extracellular matrix extending throughout the biofilm (5, 84, 131, 134, 166, 173, 199).”
§ Candida albicans Biofilms and Human Disease > IN VITRO AND IN VIVO DEVELOPMENT OF C. ALBICANS BIOFILMS
What the sources say
plain- Asserted: passes — The authors put it forward as true.
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A result the paper reports from its own data — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“Based on the current literature, we count 50 transcriptional regulators (Table 1) and 101 nonregulatory genes (Table 2) that have functionally validated roles in biofilm formation.”
§ Candida albicans Biofilms and Human Disease > GENETICS OF C. ALBICANS BIOFILM FORMATION
plain- Asserted: passes — The authors put it forward as true.
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In the paper
“Vaginal mucosal mouse models, both in vivo (live mice inoculated with C. albicans on the vaginal mucosa) and ex vivo (vaginas excised from euthanized mice and then inoculated with C. albicans in tissue culture plates), show similar biofilm architectures, with clear yeast, hyphae, and extracellular matrix evident throughout the biofilms formed on top of the mucosal layers (72).”
§ Candida albicans Biofilms and Human Disease > IN VITRO AND IN VIVO DEVELOPMENT OF C. ALBICANS BIOFILMS
What the sources say
- Not checked yetcites [72] source
boostedcorrelational- Asserted: passes — The authors put it forward as true.
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In the paper
“For example, Candida biofilms obtained from patients with denture stomatitis and from patients with infected intravascular catheters confirm the presence of yeast, hyphae and extracellular matrix (114, 162).”
§ Candida albicans Biofilms and Human Disease > IN VITRO AND IN VIVO DEVELOPMENT OF C. ALBICANS BIOFILMS
What the sources say
plain- Asserted: passes — The authors put it forward as true.
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In the paper
“Deletion of RLM1 causes a reduction in matrix levels (136), whereas deletion of ZAP1 leads to an increase in the accumulation of extracellular matrix material, probably by upregulating the glucoamylases Gca1 and Gca2 (144).”
§ Candida albicans Biofilms and Human Disease > GENETICS OF C. ALBICANS BIOFILM FORMATION > Extracellular Matrix
What the sources say
- Not checked yetcites [136] source
plaindirect- Asserted: passes — The authors put it forward as true.
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In the paper
“Of these 30, 4 (Bcr1, Ace2, Snf5, and Arg81) were also required for biofilm formation under common conditions (shaking in microtiter plates) for in vitro biofilm formation.”
§ Candida albicans Biofilms and Human Disease > GENETICS OF C. ALBICANS BIOFILM FORMATION > Adherence
What the sources say
- Not checked yetcites [57] source
plaindirect- Asserted: passes — The authors put it forward as true.
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- Load-bearing: unclear — It is unclear whether the paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
- It may state two separate claims as one. 38
- Probably not a substantial claim. 20
In the paper
“Deletion of RLM1 causes a reduction in matrix levels (136), whereas deletion of ZAP1 leads to an increase in the accumulation of extracellular matrix material, probably by upregulating the glucoamylases Gca1 and Gca2 (144).”
§ Candida albicans Biofilms and Human Disease > GENETICS OF C. ALBICANS BIOFILM FORMATION > Extracellular Matrix
What the sources say
- Not checked yetcites [144] source
hedgeddirect- Asserted: passes — The authors put it forward as true.
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It may state two separate claims as one.
In the paper
“Cells are dispersed continuously throughout biofilm formation, and they are thought to be primarily (if not exclusively) in the round yeast form (194).”
§ Candida albicans Biofilms and Human Disease > GENETICS OF C. ALBICANS BIOFILM FORMATION > Dispersal
What the sources say
- Not checked yetcites [194] source
hedged- Asserted: passes — The authors put it forward as true.
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In the paper
“In addition to the 6 master transcriptional regulators discussed above, another 44 transcriptional regulators have been identified; if any of these is deleted, some aspect of C. albicans biofilm formation is affected under at least one condition (Table 1; revised from 61 and 18).”
§ Candida albicans Biofilms and Human Disease > GENETICS OF C. ALBICANS BIOFILM FORMATION > Regulatory Control of C. albicans Biofilm Development
plaindirect- Asserted: passes — The authors put it forward as true.
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It may state two separate claims as one.
In the paper
“These include (relative to planktonic cells) increased adherence properties, the ability to form biofilms more efficiently, and enhanced virulence in mouse models of infection (194).”
§ Candida albicans Biofilms and Human Disease > GENETICS OF C. ALBICANS BIOFILM FORMATION > Dispersal
What the sources say
- Not checked yetcites [194] source
plaindirect- Asserted: passes — The authors put it forward as true.
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- Load-bearing: unclear — It is unclear whether the paper’s conclusions depend on it.
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It may state two separate claims as one.
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In the paper
“Two transcriptional regulators of C. albicans biofilm dispersal have been identified, Nrg1 and Ume6; transcriptional overexpression of either regulator increased the number of dispersed cells actively released from the biofilm (194, 195).”
§ Candida albicans Biofilms and Human Disease > GENETICS OF C. ALBICANS BIOFILM FORMATION > Dispersal
plaindirect- Asserted: passes — The authors put it forward as true.
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It may state two separate claims as one.
In the paper
“Indeed, the master regulator Bcr1 is not needed for hyphal formation per se, but it is needed for the hyphae to adhere to one another in biofilms (141).”
§ Candida albicans Biofilms and Human Disease > GENETICS OF C. ALBICANS BIOFILM FORMATION > Hyphae
What the sources say
- Not checked yetcites [141] source
plaindirect- Asserted: passes — The authors put it forward as true.
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- Load-bearing: unclear — It is unclear whether the paper’s conclusions depend on it.
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It may state two separate claims as one.
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In the paper
“The cell wall protein Ywp1 is also important for biofilm dispersal, as deletion of YWP1 leads to decreased biofilm dispersal and increased biofilm adhesiveness (69, 70).”
§ Candida albicans Biofilms and Human Disease > GENETICS OF C. ALBICANS BIOFILM FORMATION > Dispersal
plaindirect- Asserted: passes — The authors put it forward as true.
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It may state two separate claims as one.
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In the paper
“In planktonic cells, these efflux pumps are typically upregulated in response to antifungal drugs; in biofilms, however, they become upregulated within the first few hours of surface contact and remain upregulated throughout biofilm development, whether or not an antifungal drug is present (119, 126, 133, 140, 156, 205).”
§ Candida albicans Biofilms and Human Disease > GENETICS OF C. ALBICANS BIOFILM FORMATION > Drug Resistance and the Extracellular Matrix
What the sources say
plaindirect- Asserted: passes — The authors put it forward as true.
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In the paper
“Finally, extracellular DNA is also a component of the matrix in C. albicans and contributes, probably indirectly, to drug resistance (117).”
§ Candida albicans Biofilms and Human Disease > GENETICS OF C. ALBICANS BIOFILM FORMATION > Drug Resistance and the Extracellular Matrix
What the sources say
- Not checked yetcites [117] source
hedgedconditional- Asserted: passes — The authors put it forward as true.
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- Load-bearing: unclear — It is unclear whether the paper’s conclusions depend on it.
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It may state two separate claims as one.
In the paper
“Indeed, mutant strains of individual Set3 complex members formed extra strong biofilms that were incapable of normal biofilm dispersal and especially recalcitrant to mechanical perturbation (139).”
§ Candida albicans Biofilms and Human Disease > GENETICS OF C. ALBICANS BIOFILM FORMATION > Dispersal
What the sources say
- Not checked yetcites [139] source
plaindirect- Asserted: passes — The authors put it forward as true.
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- Load-bearing: unclear — It is unclear whether the paper’s conclusions depend on it.
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It may state two separate claims as one.
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In the paper
“Thus, treatment of biofilms with β-1,3-glucanase increases the susceptibility of biofilms to fluconazole (130), and addition of exogenous β-1,3-glucans increases the tolerance of planktonic cells to fluconazole (122).”
§ Candida albicans Biofilms and Human Disease > GENETICS OF C. ALBICANS BIOFILM FORMATION > Drug Resistance and the Extracellular Matrix
plaindirect- Asserted: passes — The authors put it forward as true.
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It may state two separate claims as one.
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In the paper
“Nrg1 and Set3 complex mutants are hyperfilamentous, consistent with their inability to generate yeast-form cells (77, 195).”
§ Candida albicans Biofilms and Human Disease > GENETICS OF C. ALBICANS BIOFILM FORMATION > Dispersal
plaindirect- Asserted: passes — The authors put it forward as true.
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- Load-bearing: unclear — It is unclear whether the paper’s conclusions depend on it.
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It may state two separate claims as one.
The paper states this without a source or data behind it here, so there is nothing to check it against. The quote shows it is in the paper.
In the paper
“Each of these species can form a biofilm in vitro, but the thickness, strength, and robustness in different environmental conditions decrease as species diverge phylogenetically away from C. albicans.”
§ Candida albicans Biofilms and Human Disease > BIOFILM FORMATION BY OTHER ASCOMYCOTA SPECIES
plaincorrelational- Asserted: passes — The authors put it forward as true.
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It may state two separate claims as one.
The paper states this without a source or data behind it here, so there is nothing to check it against. The quote shows it is in the paper.
In the paper
“In individuals with healthy immune systems, C. albicans is often harmless, kept in balance with other members of the local microbiota.”
§ Candida albicans Biofilms and Human Disease > INTRODUCTION
plain- Asserted: passes — The authors put it forward as true.
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Probably not a substantial claim. The paper’s conclusions may not depend on it.
The paper states this without a source or data behind it here, so there is nothing to check it against. The quote shows it is in the paper.
In the paper
“C. albicans biofilms are intrinsically resistant to conventional antifungal therapeutics, the host immune system, and other environmental perturbations, making biofilm-based infections a significant clinical challenge.”
§ Candida albicans Biofilms and Human Disease > Abstract
plaindirect- Asserted: passes — The authors put it forward as true.
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Probably not a substantial claim. The paper may give no reason why it holds. It may state two separate claims as one.
None of the sources cited for this claim have been checked yet.
In the paper
“Candida infections are especially serious in immunocompromised individuals (such as those with AIDS or those undergoing anticancer or immunosuppression therapies) and healthy people with implanted medical devices (96, 201).”
§ Candida albicans Biofilms and Human Disease > INTRODUCTION
What the sources say
- Not checked yetcites [96] Kullberg BJ, 2002, Eur. J. Med. Res.
- Not checked yetcites [201] source
plaincorrelational- Asserted: passes — The authors put it forward as true.
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In the paper
“In this review, we focus on C. albicans biofilms, which colonize implanted medical devices and mucosal surfaces, from which they can seed systemic infections in humans (46, 47, 64, 89, 97, 161, 172).”
§ Candida albicans Biofilms and Human Disease > INTRODUCTION
What the sources say
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
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It may state two separate claims as one.
The paper states this without a source or data behind it here, so there is nothing to check it against. The quote shows it is in the paper.
In the paper
“Biofilms are notorious for forming on implanted medical devices, including catheters, pacemakers, dentures, and prosthetic joints, which provide a surface and sanctuary for biofilm growth.”
§ Candida albicans Biofilms and Human Disease > Abstract
boosteddirect- Asserted: passes — The authors put it forward as true.
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- Based: fails — The paper may give no reason why it holds.
Probably not a substantial claim. The paper may give no reason why it holds.
None of the sources cited for this claim have been checked yet.
In the paper
“These infections range from superficial mucosal and dermal infections, such as thrush, vaginal yeast infections, and diaper rash, to hematogenously disseminated infection with sizable mortality rates (approaching 40% in some cases) (21, 148, 202).”
§ Candida albicans Biofilms and Human Disease > INTRODUCTION
What the sources say
plain- Asserted: passes — The authors put it forward as true.
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Probably not a substantial claim. The paper’s conclusions may not depend on it.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“It is now recognized that, for many microorganisms, the biofilm state is probably more relevant to their “natural” settings than to suspension cultures.”
§ Candida albicans Biofilms and Human Disease > INTRODUCTION
hedgedcorrelational- Asserted: passes — The authors put it forward as true.
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Probably not a substantial claim. The paper’s conclusions may not depend on it. The paper may give no reason why it holds.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“In the laboratory, C. albicans biofilms can develop on several substrates and in many types of media, indicating an inherent robustness of biofilm development to changes in external conditions.”
§ Candida albicans Biofilms and Human Disease > IN VITRO AND IN VIVO DEVELOPMENT OF C. ALBICANS BIOFILMS
hedgeddirect- Asserted: passes — The authors put it forward as true.
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Probably not a substantial claim. The paper’s conclusions may not depend on it. The paper may give no reason why it holds. It may state two separate claims as one.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“Once it forms on an implanted medical device, a Candida biofilm has the potential to seed disseminated bloodstream infections and to lead to invasive systemic infections of tissues and organs.”
§ Candida albicans Biofilms and Human Disease > INTRODUCTION
hedgedconditional- Asserted: passes — The authors put it forward as true.
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It may state two separate claims as one.
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In the paper
“Urinary and central venous catheters, pacemakers, mechanical heart valves, joint prostheses, contact lenses, and dentures are all susceptible to C. albicans biofilms (22, 45, 90, 175).”
§ Candida albicans Biofilms and Human Disease > INTRODUCTION
What the sources say
plaincorrelational- Asserted: passes — The authors put it forward as true.
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- Load-bearing: unclear — It is unclear whether the paper’s conclusions depend on it.
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It may state two separate claims as one.
None of the sources cited for this claim have been checked yet.
In the paper
“Accounting for 15% of all hospital-acquired sepsis cases, species within the CTG clade (predominantly C. albicans, but including several closely related species as well) are the fourth most frequent cause of bloodstream infections in clinical settings and are the predominant fungal species isolated from medical device infections (43, 152, 202, 204).”
§ Candida albicans Biofilms and Human Disease > INTRODUCTION
What the sources say
plain- Asserted: passes — The authors put it forward as true.
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Probably not a substantial claim. The paper’s conclusions may not depend on it. It may state two separate claims as one.
The paper states this without a source or data behind it here, so there is nothing to check it against. The quote shows it is in the paper.
In the paper
“C. albicans is distinguished from many other fungal species by its ability to form both yeast cells and hyphae under many different environmental conditions.”
§ Candida albicans Biofilms and Human Disease > GENETICS OF C. ALBICANS BIOFILM FORMATION > Hyphae
plain- Asserted: passes — The authors put it forward as true.
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- Load-bearing: fails — The paper’s conclusions may not depend on it.
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Probably not a substantial claim. The paper’s conclusions may not depend on it.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“It is clear that biofilm formation by C. albicans can occur over a broad range of conditions and that the genetic requirements likely vary from one condition to the next.”
§ Candida albicans Biofilms and Human Disease > GENETICS OF C. ALBICANS BIOFILM FORMATION > Adherence
hedgedcorrelational- Asserted: passes — The authors put it forward as true.
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It may state two separate claims as one.
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In the paper
“Although they readily form in planktonic cultures, hyphae are an important structural component of C. albicans biofilms; thus, it is not surprising that genes required for hyphal growth in suspension cultures are also necessary for proper biofilm formation.”
§ Candida albicans Biofilms and Human Disease > GENETICS OF C. ALBICANS BIOFILM FORMATION > Hyphae
What the sources say
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
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It may state two separate claims as one.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“Orthology mapping indicates that the entire set of target genes is significantly enriched for “young” genes, suggesting that the ability of C. albicans to form biofilms evolved relatively recently with respect to evolutionary timescales.”
§ Candida albicans Biofilms and Human Disease > GENETICS OF C. ALBICANS BIOFILM FORMATION > Regulatory Control of C. albicans Biofilm Development
hedgedcorrelational- Asserted: passes — The authors put it forward as true.
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It may state two separate claims as one.
In the paper
“A minor subset of metabolically dormant yeast cells that stochastically arise as phenotypical variants within biofilms, persister cells are extremely resistant to antifungal drugs (100).”
§ Candida albicans Biofilms and Human Disease > GENETICS OF C. ALBICANS BIOFILM FORMATION > Persister Cells
What the sources say
- Not checked yetcites [100] source
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: unclear — It is unclear whether the paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
In the paper
“Most of these were predicted to be enzymes, including hydrolyzing enzymes, suggesting that the matrix may play an active role in breaking down biopolymers.”
§ Candida albicans Biofilms and Human Disease > GENETICS OF C. ALBICANS BIOFILM FORMATION > Drug Resistance and the Extracellular Matrix
What the sources say
- Not checked yetcites [209] source
hedgedconditional- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
None of the sources cited for this claim have been checked yet.
In the paper
“Thus, although β-1,3-glucans contribute to drug resistance and are the major cell wall polysaccharide (130), we now know that they constitute only a small fraction of the total polysaccharides present, implying that the matrix is not constructed simply of released cell wall constituents.”
§ Candida albicans Biofilms and Human Disease > GENETICS OF C. ALBICANS BIOFILM FORMATION > Drug Resistance and the Extracellular Matrix
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
None of the sources cited for this claim have been checked yet.
In the paper
“Secreted extracellular matrix also contributes to biofilm drug resistance, both acting as a physical barrier to drug penetration and directly contributing to the overall structural integrity of the biofilm (2, 11, 130, 135).”
§ Candida albicans Biofilms and Human Disease > GENETICS OF C. ALBICANS BIOFILM FORMATION > Drug Resistance and the Extracellular Matrix
What the sources say
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“This resistance is due to the upregulation of efflux pumps, the presence of the extracellular matrix, and the presence of recalcitrant persister cells.”
§ Candida albicans Biofilms and Human Disease > GENETICS OF C. ALBICANS BIOFILM FORMATION > Drug Resistance and the Extracellular Matrix
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
None of the sources cited for this claim have been checked yet.
In the paper
“Blastoschizomyces capitatus is another rare cause of infection in humans, although more cases have been emerging in highly immunocompromised patients in ICUs.”
§ Candida albicans Biofilms and Human Disease > BIOFILM FORMATION BY OTHER ASCOMYCOTA SPECIES
What the sources say
plaincorrelational- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: fails — The paper’s conclusions may not depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim. The paper’s conclusions may not depend on it. It may state two separate claims as one.
None of the sources cited for this claim have been checked yet.
In the paper
“Along with H. capsulatum, Coccidioides immitis, Paracoccidioides brasiliensis, and Blastomyces dermatitidis are all sources of mycoses in immunocompromised individuals and have also caused endemic outbreaks in specific geographical regions (62, 113).”
§ Candida albicans Biofilms and Human Disease > BIOFILM FORMATION BY OTHER ASCOMYCOTA SPECIES
plaincorrelational- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: fails — The paper’s conclusions may not depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim. The paper’s conclusions may not depend on it. It may state two separate claims as one.
In the paper
“suggesting that H. capsulatum may exist in biofilms in vivo (154).”
§ Candida albicans Biofilms and Human Disease > BIOFILM FORMATION BY OTHER ASCOMYCOTA SPECIES
What the sources say
- Not checked yetcites [154] source
hedgedcorrelational- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: fails — The paper’s conclusions may not depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim. The paper’s conclusions may not depend on it.
In the paper
“There is at least one report of H. capsulatum forming biofilms consisting of dense clusters of yeast-form cells in vitro”
§ Candida albicans Biofilms and Human Disease > BIOFILM FORMATION BY OTHER ASCOMYCOTA SPECIES
What the sources say
- Not checked yetcites [154] source
hedged- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: fails — The paper’s conclusions may not depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim. The paper’s conclusions may not depend on it.
None of the sources cited for this claim have been checked yet.
In the paper
“and it is able to form a thin biofilm consisting of round, budding yeast-form cells and pseudohyphal cells in vitro (26, 50, 128, 164).”
§ Candida albicans Biofilms and Human Disease > BIOFILM FORMATION BY OTHER ASCOMYCOTA SPECIES
What the sources say
plain- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: fails — The paper’s conclusions may not depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim. The paper’s conclusions may not depend on it.
In the paper
“Aspergillus fumigatus is a common invasive colonizer of the respiratory tract in individuals with cystic fibrosis or other conditions that compromise the immune system (176).”
§ Candida albicans Biofilms and Human Disease > BIOFILM FORMATION BY OTHER ASCOMYCOTA SPECIES
What the sources say
- Not checked yetcites [176] source
plaincorrelational- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: fails — The paper’s conclusions may not depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim. The paper’s conclusions may not depend on it.
In the paper
“Aspergillomas, or dense hyphal balls, may also form as A. fumigatus continues to proliferate in the respiratory tract; these structures share many architectural characteristics with biofilms formed by other fungal species (159).”
§ Candida albicans Biofilms and Human Disease > BIOFILM FORMATION BY OTHER ASCOMYCOTA SPECIES
What the sources say
- Not checked yetcites [159] source
hedgedconditional- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: unclear — It is unclear whether the paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
The paper states this without a source or data behind it here, so there is nothing to check it against. The quote shows it is in the paper.
In the paper
“Outside the CTG clade, numerous fungal species have been reported to form biofilms in vitro and in vivo.”
§ Candida albicans Biofilms and Human Disease > BIOFILM FORMATION BY OTHER ASCOMYCOTA SPECIES
plain- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: fails — The paper’s conclusions may not depend on it.
- Based: fails — The paper may give no reason why it holds.
Probably not a substantial claim. The paper’s conclusions may not depend on it. The paper may give no reason why it holds.
None of the sources cited for this claim have been checked yet.
In the paper
“Aspergillus is a fungal pathogen of great medical importance that can form biofilms both in vitro and in vivo (125, 159).”
§ Candida albicans Biofilms and Human Disease > BIOFILM FORMATION BY OTHER ASCOMYCOTA SPECIES
plain- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: fails — The paper’s conclusions may not depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim. The paper’s conclusions may not depend on it. It may state two separate claims as one.
In the paper
“Histoplasma capsulatum is another opportunistic pathogen that infects the respiratory system of mammals, causing histoplasmosis (85).”
§ Candida albicans Biofilms and Human Disease > BIOFILM FORMATION BY OTHER ASCOMYCOTA SPECIES
What the sources say
- Not checked yetcites [85] source
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: fails — The paper’s conclusions may not depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim. The paper’s conclusions may not depend on it. It may state two separate claims as one.
None of the sources cited for this claim have been checked yet.
In the paper
“The closely related species Candida dubliniensis, Candida parapsilosis, Candida tropicalis, and Candida krusei have all been implicated in biofilm-associated infections (73, 157, 187).”
§ Candida albicans Biofilms and Human Disease > BIOFILM FORMATION BY OTHER ASCOMYCOTA SPECIES
What the sources say
plaincorrelational- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: unclear — It is unclear whether the paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
None of the sources cited for this claim have been checked yet.
In the paper
“In a few case studies, however, S. cerevisiae has been implicated in catheter-associated infections with mixed-species biofilms in patients in intensive care units (ICUs)”
§ Candida albicans Biofilms and Human Disease > BIOFILM FORMATION BY OTHER ASCOMYCOTA SPECIES
What the sources say
hedgedcorrelational- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: fails — The paper’s conclusions may not depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim. The paper’s conclusions may not depend on it.
In the paper
“Although rare, C. immitis has been reported to form biofilms on medical devices (40).”
§ Candida albicans Biofilms and Human Disease > BIOFILM FORMATION BY OTHER ASCOMYCOTA SPECIES
What the sources say
- Not checked yetcites [40] source
hedged- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: fails — The paper’s conclusions may not depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim. The paper’s conclusions may not depend on it.
A conclusion the paper draws from its own results — it cites no other source for it, so there is none to check. The quote shows it is in the paper.
In the paper
“From these studies, we have learned that C. albicans and bacteria can interact with each other by secretion of signaling molecules that influence the behavior of one species toward the other, by direct cell-cell physical contact, and by alterations of the local environment that influence the other species (e.g., alterations in pH and oxygen concentrations).”
§ Candida albicans Biofilms and Human Disease > MULTISPECIES BIOFILMS WITH C. ALBICANS
plaindirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
In the paper
“Recent work in vitro has shown that F. solani, A. implicatum, and C. sphaerospermum can all form biofilms under laboratory conditions (211).”
§ Candida albicans Biofilms and Human Disease > BIOFILM FORMATION BY OTHER ASCOMYCOTA SPECIES
What the sources say
- Not checked yetcites [211] source
plain- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: unclear — It is unclear whether the paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
None of the sources cited for this claim have been checked yet.
In the paper
“and it is widely believed that antifungal drugs with new mechanisms of action are needed (4, 18, 79).”
§ Candida albicans Biofilms and Human Disease > PERSPECTIVES FOR DEVELOPING THERAPEUTICS FOR C. ALBICANS BIOFILM INFECTIONS
What the sources say
plain- Asserted: passes — The authors put it forward as true.
- Checkable: fails — It may be too vague to check against evidence.
- Load-bearing: fails — The paper’s conclusions may not depend on it.
- Based: passes — The paper gives a reason why it holds.
Probably not a substantial claim. It may be too vague to check against evidence. The paper’s conclusions may not depend on it.
None of the sources cited for this claim have been checked yet.
In the paper
“C. albicans also interacts with several bacterial species found in the gut, such as Enterococcus and Escherichia species; those found in the vagina, such as Lactobacillus species; and those found in the lungs of patients with cystic fibrosis, such as Pseudomonas aeruginosa (13, 34, 54, 78).”
§ Candida albicans Biofilms and Human Disease > MULTISPECIES BIOFILMS WITH C. ALBICANS
What the sources say
plaincorrelational- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: unclear — It is unclear whether the paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
In the paper
“Moreover, when cultured together with C. albicans in ambient oxygen, these anaerobes could induce the latter to form mini-biofilms, which, in turn, could protect the bacteria and allow them to proliferate under otherwise toxic conditions.”
§ Candida albicans Biofilms and Human Disease > MULTISPECIES BIOFILMS WITH C. ALBICANS
What the sources say
- Not checked yetcites [60] source
hedgeddirect- Asserted: passes — The authors put it forward as true.
- Checkable: passes — It can be checked against evidence.
- Load-bearing: passes — The paper’s conclusions depend on it.
- Based: passes — The paper gives a reason why it holds.
It may state two separate claims as one.
The paper relies on this without support.
Virtually nothing is known about the interactions among Candida species in polymicrobial biofilms.
“We know virtually nothing about the interactions among these Candida species in polymicrobial biofilms.”
The paper relies on this without support.
No biofilm-specific drugs exist today for Candida albicans or any microbe.
“No biofilm-specific drugs exist today for C. albicans (or any microbe), making treatment of biofilm-based infections particularly problematic.”
The paper relies on this without support.
Candida albicans is the fungal pathogen most frequently isolated from human infections.
“Although C. albicans is the fungal pathogen most frequently isolated from human infections”
The paper relies on this without support.
Microbial infections are often thought of, and treated, as if a single microbial species were acting alone.
“Microbial infections are often thought of, and treated, as if a single microbial species were acting alone.”
The paper relies on this without support.
Keratitis is generally thought of as a biofilm infection and has been associated with the use of contact lenses.
“Keratitis is generally thought of as a biofilm infection and has been associated with the use of contact lenses.”
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?
The cited source says what the claim attributes to it.
In the citing paper
“Once established, these biofilms exhibit a high tolerance to antifungal drugs, posing a substantial challenge in clinical treatment [54,55,56,57].”
From Disruption of the UPC2 Gene Enhances Fluconazole Antifungal Activity by Inhibiting HAC1 mRNA Splicing in Candida albicans (2026) · cites this paper as [56]
In the citing paper
“A well-documented virulence factor of Candida species is the notable ability to form adherent structures, known as biofilms, on surfaces of medical devices [49–51].”
From Aprepitant, an antiemetic agent, interferes with metal ion homeostasis of Candida auris and displays potent synergistic interactions with azole drugs (2020) · cites this paper as [50]
Citations
Cites
Items 201–215 of 215
Candida albicans PEP12 Is Required for Biofilm Integrity and In Vivo Virulence
Palanisamy, S. K. A., Ramírez, M. A., Lorenz, M., & Lee, S. A. (2009) · Eukaryotic Cell · doi:10.1128/ec.00295-09 · Cited by 33
Seddiki, S. M. L., Boucherit-otmani, Boucherit, Badsi-amir, Taleb, & Kunkel (2013) · International Journal of General Medicine · doi:10.2147/ijgm.s38065 · Cited by 27
In vitro analysis of flufenamic acid activity against Candida albicans biofilms
Chavez-Dozal, A. A., Jahng, M., Rane, H. S., Asare, K., Kulkarny, V. V., Bernardo, S. M., & Lee, S. A. (2014) · International Journal of Antimicrobial Agents · doi:10.1016/j.ijantimicag.2013.08.018 · Cited by 26
Bioluminescence Imaging of Fungal Biofilm Development in Live Animals
Velde, G. V., Kucharíková, S., Dijck, P. V., & Himmelreich, U. (2013) · Methods in molecular biology · doi:10.1007/978-1-62703-718-1_13 · Cited by 25
Sen, M., Shah, B., Rakshit, S., Singh, V., Padmanabhan, B., Ponnusamy, M., Pari, K., Vishwakarma, R. A., Nandi, D., & Sadhale, P. P. (2011) · PLoS Pathogens · doi:10.1371/journal.ppat.1002384 · Cited by 25
Pga26 mediates filamentation and biofilm formation and is required for virulence in Candida albicans
Laforet, L., Moreno, I., Sánchez-Fresneda, R., Martinez‐Esparza, M., Martínez, J. P., Argüelles, J. C., de Groot, P. W. J., & Valentı́n, E. (2011) · FEMS Yeast Research · doi:10.1111/j.1567-1364.2011.00727.x · Cited by 24
Rouabhia, M., Semlali, A., Chandra, J., Mukherjee, P. K., Chmielewski, W., & Ghannoum, M. A. (2012) · Mediators of Inflammation · doi:10.1155/2012/398207 · Cited by 19
A proteomic analysis of secretory proteins of a pre-vacuolar mutant of Candida albicans
Thomas, D. P., Lopez-Ribot, J. L., & Lee, S. A. (2009) · Journal of Proteomics · doi:10.1016/j.jprot.2009.10.003 · Cited by 15
Pietrella, D., Enjalbert, B., Zeidler, U., Znaidi, S., Rachini, A., Vecchiarelli, A., & d’Enfert, C. (2012) · Methods in molecular biology · doi:10.1007/978-1-61779-539-8_39 · Cited by 14
The role of Mss11 in Candida albicans biofilm formation
Tsai, P.-W., Chen, Y.-T., Yang, C., Chen, H.-F., Tan, T.-S., Lin, T.-W., Hsieh, W., & Lan, C. (2014) · Molecular Genetics and Genomics · doi:10.1007/s00438-014-0846-0 · Cited by 13
Experimental candidiasis after thermal injury
Fader, R., Nunez, D., Unbehagen, J., & Linares, H. A. (1985) · Infection and Immunity · doi:10.1128/iai.49.3.780-784.1985 · Cited by 12
Daher, J., Koussa, J., Younes, S., & Khalaf, R. A. (2011) · Interdisciplinary Perspectives on Infectious Diseases · doi:10.1155/2011/504280 · Cited by 12
Drug susceptibility of matrix‐encapsulated Candida albicans nano‐biofilms
Srinivasan, A., Gupta, C. M., Agrawal, C. M., Leung, K. P., Lopez‐Ribot, J. L., & Ramasubramanian, A. K. (2014) · Biotechnology and Bioengineering · doi:10.1002/bit.25120 · Cited by 11
Melo, A. S. D. A., Padovan, A. C. B., Serafim, R. C., Puzer, L., Carmona, A. K., Juliano, L., Brunstein, A., & Briones, M. R. S. (2006) · PubMed · Cited by 10
Sun, X., Lu, H., Jiang, Y., & Cao, Y. (2013) · Current Microbiology · doi:10.1007/s00284-013-0366-x · Cited by 5
Research works printed but not linked: 5
- Dominic RM, 2007, Kathmandu Univ. Med. J., 5, 431
- Douglas LJ, 2002, Rev. Iberoamericana Micol., 19, 139
- Kolter R, 2010, Int. Microbiol., 13, 1
- Kullberg BJ, 2002, Eur. J. Med. Res., 7, 183
- Melo AS, 2006, Genet. Mol. Res., 5, 664
Cited by
Sorted by citations
Candida albicans—The Virulence Factors and Clinical Manifestations of Infection
Talapko, J., Juzbašić, M., Matijević, T., Pustijanac, E., Bekić, S., Kotris, I., & Škrlec, I. (2021) · Journal of Fungi · doi:10.3390/jof7020079 · Cited by 794
“albicans cells phagocytosed by host immune cells, direct damage of mucosal barriers, and spread from fungal biofilms [49,91].”
BackgroundCandida albicans biofilms: development, regulation, and molecular mechanisms
Gulati, M., & Nobile, C. J. (2016) · Microbes and Infection · doi:10.1016/j.micinf.2016.01.002 · Cited by 738
Self-citationVaginal microbiota and the potential of Lactobacillus derivatives in maintaining vaginal health
Chee, W. J. Y., Chew, S. Y., & Than, L. T. L. (2020) · Microbial Cell Factories · doi:10.1186/s12934-020-01464-4 · Cited by 761
Development and regulation of single- and multi-species Candida albicans biofilms
Lohse, M. B., Gulati, M., Johnson, A. D., & Nobile, C. J. (2018) · Nature Reviews Microbiology · doi:10.1038/nrmicro.2017.107 · Cited by 643
Self-citationCandida auris: Epidemiology, biology, antifungal resistance, and virulence
Du, H., Bing, J., Hu, T., Ennis, C. L., Nobile, C. J., & Huang, G. (2020) · PLOS Pathogens · doi:10.1371/journal.ppat.1008921 · Cited by 609
Self-citation
Metadata
- Authors
- Nobile, C. J., & Johnson, A. D.
- Affiliations
- University of California, Merced (US)
- University of California, San Francisco (US)
- Venue
- Annual Review of Microbiology
- Publisher
- Annual Reviews
- Published
- 2015-10-15
- Volume
- 69
- Issue
- 1
- Pages
- 71-92
- ISSN
- 0066-4227 (ISSN-L)
- 1545-3251
- PMC
- PMC4930275
- OpenAlex
- W2170764510
- Semantic Scholar
- 5e945268f6f8fd4d66044feb13a7f07654261596
- Cited by
- 1,057
- References
- 215
- Citations by year
- 2026: 77 · 2025: 106 · 2024: 130 · 2023: 88 · 2022: 138 · 2021: 135 · 2020: 117 · 2019: 91 · 2018: 76 · 2017: 68 · 2016: 28 · 2015: 2
- FWCI
- 8.81
- Type
- journal article
- Language
- en
- Open access
- bronze
- doi.org
- Locations
- Annual Review of Microbiology (publishedVersion)
- PubMed (publishedVersion)
- PubMed Central (acceptedVersion)
- eScholarship (California Digital Library) (submittedVersion)
- Funders
- National Institutes of Health
- National Institute of Allergy and Infectious Diseases (R00AI100896, R01AI083311, T32AI060537)
- Topics
- Health Sciences › Medicine › Infectious Diseases › Antifungal resistance and susceptibility
- Life Sciences › Biochemistry, Genetics and Molecular Biology › Molecular Biology › Bacterial biofilms and quorum sensing
- Life Sciences › Biochemistry, Genetics and Molecular Biology › Molecular Biology › Fungal and yeast genetics research
- Keywords
- Candida albicans, biofilm, candidiasis, fungal infection, human microbiota, antifungal resistance, host immune response, biomedical implants, biofilm-associated infections, mucosal candidiasis, systemic candidiasis, fungal communities
- MeSH terms
- Animals, Candida albicans, Candidiasis, Cell Adhesion, Fungi, Humans, Mycoses, Biofilms
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.
A language model drew the claims from the paper on 2026-10-09, under version 3 of the claim definition (model deepseek-v4.1-flash, prompt extract-v3).
A second model labelled each claim on 2026-10-09 (model jev-1.13-20260917, definition revision gold-guideline-v0.2).
Cited sources are checked with a language model’s help (gpt-6-sol).
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.
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