Architecture of the dynamic fungal cell wall
Open accessAbstract
The fungal cell wall is essential for growth and survival, and is a key target for antifungal drugs and the immune system. The cell wall must be robust but flexible, protective and shielding yet porous to nutrients and membrane vesicles and receptive to exogenous signals. Most fungi have a common inner wall skeleton of chitin and β-glucans that functions as a flexible viscoelastic frame to which a more diverse set of outer cell wall polymers and glycosylated proteins are attached. Whereas the inner wall largely determines shape and strength, the outer wall confers properties of hydrophobicity, adhesiveness, and chemical and immunological heterogeneity. The spatial organization and dynamic regulation of the wall in response to prevailing growth conditions enable fungi to thrive within changing, diverse and often hostile environments. Understanding this architecture provides opportunities to develop diagnostics and drugs to combat life-threatening fungal infections. In this Review, Gow and Lenardon describe how fungal cell walls are organized, focusing on the underlying architectural and mechanical principles that are required to deliver differing and bespoke biochemical and biophysical attributes.
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C. auris cell walls contain a higher abundance of N-linked mannans compared to O-linked mannans.
“The cell walls of C. auris, like C. albicans, contain a higher abundance of N-linked mannans compared to O-linked mannans (23).”
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 [23]
C. albicans has a well-characterized cell wall composed of an inner layer of chitin, β-1,3-glucan, and β-1,6-glucan, and an outer layer composed of proteins modified with N- and O-linked mannans and phosphomannans.
“C. albicans has a well-characterized cell wall composed of an inner layer of chitin, β−1,3-glucan, and β−1,6-glucan, and an outer layer composed of proteins modified with N- and O-linked mannans and phosphomannans, called mannoproteins34,35.”
From Distinct echinocandin responses of Candida albicans and Candida auris cell walls revealed by solid-state NMR (2025) · cites this paper as [35]
These α-1,2-mannoses can be connected to the α-1,6-mannan backbone, as well as to β-1,2-mannose, α-1,3-mannose, and phosphate groups found in the sidechains.
“These α-1,2-mannoses can be connected to the α-1,6-mannan backbone, as well as to β-1,2-mannose, α-1,3-mannose, and phosphate groups found in the sidechains27,35.”
From Distinct echinocandin responses of Candida albicans and Candida auris cell walls revealed by solid-state NMR (2025) · cites this paper as [35]
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“Most of this carbohydrate is found in branched polymers of glucose (β-glucans), unbranched linear polymers of β-1,4-N-acetylglucosa mine (chitin), and polymers of mannose (mannans) (58, 59).”
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“The fungal cell wall, an intricate and dynamic composite of polysaccharides and proteins positioned outside the plasma membrane, serves as the frontline in the interaction between fungi and their environment [1,2].”
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Marine-Derived Metabolites Act as Promising Antifungal Agents
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“The outer layer of the cell wall is formed by attaching N -mannan structures to CWPs, while short-chain O -mannan are also attached to CWPs [26] (Figure 1).”
BackgroundBreaking down the wall: Solid-state NMR illuminates how fungi build and remodel diverse cell walls
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“23 20 This similarity highlights both the robustness of the methodology and the broad applicability of this biovolume-to-carbon conversion factor across diverse ecosystems, reinforcing its potential as a reliable tool for estimating fungal biomass in various environments.”
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Strong and tough chitin hydrogel constructed by dehydration and rehydration strategy
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“The outer wall of fungal hyphae covered by the N-mannans of glycoproteins, especially the cell wall of Candida albicans (45), has been proposed to function as a b-glucan barrier to protect the inner cell walls, where GPI-anchored cell proteins and glycoproteins are located.”
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“The biomass of fungal residues mainly contains polysaccharides, which account for 80–90% of the total cell wall (Baldrian et al., 2013; Gow and Lenardon, 2022).”
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Roles of α-1,3-glucosyltransferase in growth and polysaccharides biosynthesis of Ganoderma lucidum
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Black yeasts in hypersaline conditions
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“…growth facilitates the exchange of water, nutrient absorption and solutes between cells, this connectivity also makes the cells susceptible to water loss due to the high surface area-to-volume ratio of hyphae, making them more susceptible to water loss (Sterflinger 1998; Gow and Lenardon 2023).”
BackgroundTop five unanswered questions in fungal cell surface research
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Self-citationCharting the solid‐state NMR signals of polysaccharides: A database‐driven roadmap
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“Glucans, chitins, chitosans, mannoproteins (MPs), and Glycosylphosphatidylinositol (GPI)-anchored proteins, which are the important constitutes of the cell walls, are also pathogen-associated molecular patterns (PAMPs) (Garcia-Rubio et al., 2019; Gow and Lenardon, 2023).”
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“28 These structural modi fi cations may create additional binding sites for nanoparticle interactions and increase cellular uptake ef fi ciency.”
Metals and the cell surface of Cryptococcus neoformans
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“Therefore, deactivating fungi is more challenging than general bacteria [33].”
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Magic-angle spinning NMR spectral editing of polysaccharides in whole cells using the DREAM scheme.
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Engineering Strategies for Fungal Cell Disruption in Biotechnological Applications
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Fungal β-1,3-glucan synthase: a review of structure, mechanism, and regulation
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“It forms the core fibrillar network of the cell wall, providing rigidity and structural strength (Lesage and Bussey 2006 , Gow and Lenardon 2023 ).”
Immunomodulatory function of chitosan is dependent on complement receptor 3
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“Unlike chitin, chitosan is always deacetylated from chitin, and is not a prevalent component in the cell wall of most fungi, although chitosan represents a significant fraction of the wall of Cryptococcus species and Zygomycetes (Lam, Upadhya et al. 2019, Gow and Lenardon, 2023).”
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“Echi-nocandins, in the process of inhibiting 1,3-β -glucan synthase, also trigger cell wall stress responses, which are characterized by an increase in chitin content and enhanced exposure of 1,3-β -glucan in Candida species [10,11].”
The phenomenon of anhydrobiosis—structural and functional changes in yeast cells
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“In addition to the β-glucan skeleton, the inner layer also contains chitin, which, together with β-glucan, is linked by covalent bonds (Edo et al. 2024) to form a network located in the lower part of the inner layer (Gow and Lenardon 2022).”
InfluentialExtracellular vesicle-mediated RNA warfare: A cross-kingdom battleground.
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“It is important to recognize that the fungal cell wall is a highly dynamic structure, continuously reshuffling its composition and nanoscale organization in response to the fungus's age, growth conditions, and environmental stressors (Gow & Lenardon, 2023).”
- Authors
- Gow, N. A. R., & Lenardon, M. D.
- Affiliations
- University of Exeter (GB)
- UNSW Sydney (AU)
- Corresponding author
- Gow, Neil A. R.
- Lenardon, Megan D.
- Venue
- Nature Reviews Microbiology
- Published
- 2022-10-20
- Volume
- 21
- Issue
- 4
- Pages
- 248-259
- ISSN
- 1740-1526 (ISSN-L)
- 1740-1534
- OpenAlex
- W4306878780
- Cited by
- 447
- References
- 138
- Citations by year
- 2026: 146 · 2025: 157 · 2024: 93 · 2023: 48 · 2022: 3
- FWCI
- 7.89
- Type
- journal article
- Language
- en
- Open access
- hybrid
- doi.org
- Licence
- Locations
- Nature Reviews Microbiology (publishedVersion, cc-by)
- PubMed (publishedVersion)
- UNSWorks (University of New South Wales, Sydney, Australia) (submittedVersion, cc-by-nc-nd)
- Funders
- Wellcome Trust (215599/Z/19/Z, 200208/A/15/Z, 101873/Z/13/Z)
- Medical Research Council (MR/N006364, MR/N006364/2, MR/J008230/1, MR/M026663/2, MR/V033417/1)
- Topics
- Life Sciences › Biochemistry, Genetics and Molecular Biology › Molecular Biology › Fungal and yeast genetics research
- Health Sciences › Medicine › Infectious Diseases › Antifungal resistance and susceptibility
- Health Sciences › Medicine › Epidemiology › Fungal Infections and Studies
- Keywords
- fungal cell wall, chitin, β-glucan, cell wall architecture, cell wall mechanics, viscoelasticity, glycosylated proteins, cell wall hydrophobicity, cell adhesion, cell wall regulation, antifungal drug targets, fungal infection
- MeSH terms
- Antifungal Agents, Cell Wall, Chitin, Fungal Proteins, Glucans, beta-Glucans
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