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HOME > J. Microbiol > Volume 63(12); 2025 > Article
Review
 Integrative perspectives on glycosylation networks in fungi and oomycetes
Heeji Moon1, Hokyoung Son1,2,3,4,*
Journal of Microbiology 2025;63(12):e2510003.
DOI: https://doi.org/10.71150/jm.2510003
Published online: December 31, 2025

1Department of Agricultural Biotechnology, Seoul National University, Seoul 08826, Republic of Korea

2Plant Health Center, Seoul National University, Seoul 08826, Republic of Korea

3Research Institute of Agriculture and Life Sciences, Seoul National University, Seoul 08826, Republic of Korea

4Plant Genomics and Breeding Institute, Seoul National University, Seoul 08826, Republic of Korea

*Correspondence Hokyoung Son hogongi7@snu.ac.kr
• Received: October 10, 2025   • Revised: November 19, 2025   • Accepted: November 24, 2025

© The Microbiological Society of Korea

This is an Open Access article distributed under the terms of the Creative Commons Attribution Non-Commercial License (http://creativecommons.org/licenses/by-nc/4.0) which permits unrestricted non-commercial use, distribution, and reproduction in any medium, provided the original work is properly cited.

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  • Pathogenic fungi pose major threats to both global food security and human health, yet the molecular basis of their virulence remains only partially understood. Beyond genetic and transcriptional control, emerging evidence highlights protein glycosylation as a key post-translational modification that governs fungal development, stress adaptation, and host interactions. Glycosylation regulates protein folding, stability, trafficking, and immune evasion, thereby shaping infection processes across diverse pathogens. While extensively studied in model organisms, our understanding of glycosylation in pathogenic fungi remains fragmented and lacks a coherent framework linking glycosylation dynamics to fungal development and pathogenicity. This review synthesizes recent advances from proteomic, transcriptomic, and glycomic studies in pathogenic fungi, focusing on interspecific variation in glycogenes and enzymes, hierarchical regulatory networks, and glycoprotein-mediated mechanisms of virulence. Finally, we outline current challenges and highlight glycosylation-targeted strategies as promising avenues for antifungal intervention.
  • Keywords: glycosylation, glycoprotein, pathogenicity, fungi, host-fungus interaction
Pathogenic fungi cause devastating diseases in both plants and humans, posing severe threats to global food security and public health. These infections not only result in major agricultural losses but also impose considerable medical and social burdens (Doehlemann et al., 2017; Rokas, 2022). Accordingly, elucidating fungal pathogenic mechanisms and identifying key determinants of virulence remain central objectives in developing effective prevention and control strategies (Brown et al., 2024; Sang and Kim, 2020; Yin et al., 2023).
In recent years, research has increasingly focused on the molecular basis of fungal pathogenicity. Whereas earlier studies primarily emphasized genetic and transcriptional regulation, growing evidence highlights the pivotal roles of post-translational modifications (PTMs) in fungal development, stress adaptation, and virulence (Leach and Brown, 2012). PTMs diversify protein function through covalent modifications, including the addition of small molecules or polypeptides and site-specific chemical alterations. In eukaryotes, PTMs regulate diverse cellular processes such as apoptosis, transcription, DNA repair, cell-cycle progression, and protein–protein interactions (Sartorelli et al., 1999; Tahir et al., 2019; Zhang et al., 2004).
Among the major PTMs, glycosylation stands out for its remarkable structural complexity. Rather than functioning as a simple on/off modification, glycosylation generates a continuum of glycoforms defined by site occupancy and fine structural variation (Ives et al., 2024). Glycan assembly begins with the transfer of oligosaccharides to specific amino-acid residues—Asn for N-linked glycans (within the Asn-X-Ser/Thr consensus motif) and Ser/Thr for O-linked glycans—followed by sequential processing within the endoplasmic reticulum (ER) and Golgi apparatus. Secretory and membrane glycoproteins, after acquiring glycans through the activity of mannosyltransferases, the oligosaccharyltransferase (OST) complex, and glycosidases, are delivered to the plasma membrane, cell wall, vacuoles, lysosomes, or the extracellular space (Helenius and Aebi, 2004). These processes collectively govern protein folding, stability, trafficking, localization, abundance, and biological activity, thereby shaping fungal physiology and pathogenicity (Jiang et al., 2025; Liu et al., 2021).
The origins of glycosylation research trace back to Emil Fischer’s synthesis of simple glycosides in the 1890s, which launched the modern field of carbohydrate chemistry (Haese et al., 2022). Since then, glycosylation has been comprehensively investigated in model organisms such as humans, mice, Arabidopsis thaliana, and yeast (Larskaya et al., 2024; Schjoldager et al., 2020; Tanner and Lehle, 1987). In contrast, studies in fungi remain fragmented, and a unifying framework linking glycosylation to fungal development and pathogenesis is still missing. In this review, we aim to bridge this gap by systematically examining the roles of protein glycosylation in fungal pathogens. We highlight three focal areas: (1) interspecific variation in glycogenes, defined as genes encoding glycosylation enzymes, transporters, and related factors, (2) hierarchical regulatory networks of glycosylation and their impact on pathogenicity, and (3) glycosylation-mediated mechanisms of virulence. Finally, we discuss outstanding challenges and propose future research directions, emphasizing glycosylation-targeted strategies as promising avenues for antifungal intervention.
The assembly of the core glycan precursor in the ER is highly conserved across eukaryotes (Wang et al., 2017). However, subsequent modifications in the Golgi apparatus diverge markedly among lineages, reflecting the extensive genetic repertoire of enzymes that tailor glycan structures. Each fungal species harbors a distinct set of genes encoding glycosyltransferases, glucosidases, and OST subunits, which collectively define the diversity of both N- and O-linked oligosaccharide structures (Deshpande et al., 2008; Moon et al., 2025). Consequently, comparative analysis of glycogenes provides critical insights into species-specific glycan architectures and their functional implications for fungal biology.
Glycosyltransferases
Glycosyltransferases are widely conserved in fungi, but their abundance and diversity vary considerably among lineages. Mannosyltransferases, for example, are far more prevalent in Saccharomyces cerevisiae than in filamentous fungi, reflecting its characteristic hypermannosylation. By contrast, filamentous fungi harbor a broader repertoire of genes encoding mannosyltransferases involved in high-mannose glycan maturation. This is supported by the presence of mannan polymerase I (M-Pol I; encoded by VAN1 and MNN9) and M-Pol II (MNN9, ANP1, MNN10, MNN11, and OCH1) subunits, which occur in varying copy numbers across lineages (Moon et al., 2025).
Species-specific glycosyltransferases also give rise to distinct glycan structures. Galactosyltransferases, for instance, are uniquely present in Magnaporthe oryzae, Neurospora crassa, Aspergillus nidulans, and Schizosaccharomyces pombe, but absent in S. cerevisiae, Cryptococcus neoformans, and Fusarium species, consistent with the occurrence of galactose-extended glycans only in these taxa (Deshpande et al., 2008; Moon et al., 2025). Likewise, the gene MGAT3, encoding β-1,4-mannosyl-glycoprotein 4-β-N-acetylglucosaminyltransferase, is enriched in filamentous fungi, where it likely contributes to the generation of hybrid or branched N-glycans uncommon in yeast-form fungi (Moon et al., 2025). Another notable example is C. neoformans. Although the genes encoding xylose transferases have not yet been fully characterized in this species, xylose-containing glycans are consistently detected in its cell wall and capsule (Park et al., 2012; Thak et al., 2020). These unusual xylose modifications are hallmarks of C. neoformans glycobiology and are thought to enhance antigenicity and promote immune evasion, further underscoring how species-specific enzymatic repertoires shape distinct glycan architectures.
Glucosidases
Glucosidases — notably α-glucosidase I and II, encoded by GLS1 and GLS2, respectively — are broadly conserved, yet their proportional representation differs. Filamentous fungi maintain robust glucosidase complements, whereas S. cerevisiae and S. pombe display comparatively reduced repertoires. This correlates with the observation that in yeast, trimming is minimal and often bypassed by hypermannosylation, whereas in filamentous fungi, glucosidase-mediated processing contributes to intermediate glycan forms (e.g., Man9GlcNAc2 → Man5GlcNAc2) (De Pourcq et al., 2010). Distinct modifications further highlight functional divergence: in Trichoderma reesei, insufficient trimming by glucosidase II leads to abnormal glycans with phosphodiester linkages, while Aspergillus fumigatus produces galactofuranose (Galf)-containing N- and O-glycans, reflecting species-specific enzyme contributions to antigenic structures (De Pourcq et al., 2010).
OST complex
Core subunits of the OST complex (Stt3, Ost1, and Wbp1) are highly conserved across fungi, reflecting their essential role in transferring Glc3Man9GlcNAc2to nascent polypeptides at the canonical N-glycosylation sequon Asn-X-Ser/Thr (X ≠ Pro). However, accessory subunits such as Ost3 and Ost6, required for maximal efficiency in S. cerevisiae, show lower sequence conservation in filamentous fungi. Moreover, homologs of Ost2, Swp1, and Ost4 are not consistently identifiable in genomes of filamentous fungi, suggesting possible functional divergence or replacement (Moon et al., 2025). These lineage-specific differences likely underlie variable occupancy of N-glycosylation sites and distinct glycoprotein repertoires. For example, A. fumigatus relies on Galf residues in its N-glycans and cell wall galactomannans, a structure absent in yeasts, which contributes to its immunogenicity and serves as a diagnostic marker for invasive aspergillosis (Onoue et al., 2018; Schmalhorst et al., 2008).
In recent years, substantial progress has been made in dissecting the roles of protein glycosylation in pathogenic fungi, including studies on glycogenes, structural characterization of glycans, and functional analyses of glycoproteins. However, these investigations have largely remained fragmented, focusing on isolated components rather than integrated pathways. As a result, our current understanding of how transcription factors (TFs), glycogenes, glycan structures, and glycoproteins are organized into coordinated regulatory mechanisms is still limited. A comprehensive network-level framework linking these hierarchical layers of glycosylation to fungal development and pathogenesis has yet to be established.
TFs regulating glycogenes
TFs are central regulators that orchestrate fungal growth, development, and virulence by controlling diverse sets of downstream genes. In pathogenic fungi, large-scale TF deletion mutant libraries have been established in Fusarium graminearum, C. neoformans, and M. oryzae, enabling systems-level analyses of TF networks (Jung et al., 2015; Lu et al., 2014; Son et al., 2011). These studies have uncovered regulatory hubs and functional redundancies across pathways related to virulence, secondary metabolism, development, and stress adaptation. Such systems-level insights provide integrated maps that reveal crosstalk among pathways and highlight potential vulnerabilities that may serve as antifungal targets.
By contrast, only a few TFs have been directly linked to glycogenes in fungi, whereas mammalian studies provide a more detailed picture of TF–glycogene regulation. For instance, ATF2 has been described as a paradigm of multifaceted transcriptional regulation in cancer biology (Watson et al., 2017). More recently, computational frameworks have been applied to predict TFs and signaling pathways that regulate glycan biosynthesis on the genome scale (Groth et al., 2021). Members of the CREB3-like family have also emerged as central regulators, responding to nutrient availability and ER–Golgi stress and acting as transcriptional hubs coupling glycosylation with cellular homeostasis (Khan and Margulies, 2019; Sampieri et al., 2019).
To date, only a handful of fungal TFs have been implicated in glycosylation regulation, and most remain poorly characterized. As summarized in Table 1, examples include Ace2 in Candida albicans (Cantero and Ernst, 2011; van Wijlick et al., 2016), which regulates O-mannosyltransferases (PMT) genes; Ftg1 in Botrytis cinerea (Yang et al., 2024), which influences ALG8; and unfolded protein response regulators, represented by the functional orthologs Hac1 (H. polymorpha), HacA (A. niger), and Hxl1 (C. neoformans), which activate a broad spectrum of N-glycogenes across yeasts and filamentous fungi (Carvalho et al., 2012; Cheon et al., 2011; Glazier et al., 2015; Moon et al., 2015). In addition, stress-responsive TFs such as PacC in Metarhizium robertsii (Huang et al., 2015) and Ste12 in Metarhizium acridum (Wei et al., 2017) have been implicated in the regulation of STT3, OST genes, and OCH1. Despite these examples, direct DNA–TF interactions have been experimentally confirmed in only a few cases, underscoring the need for comprehensive mapping of TF–glycogene regulatory networks in pathogenic fungi.
Glycogenes and glycan structures
While transcriptional regulators define the upstream control, the functional consequences of glycosylation are largely determined by the glycogenes themselves and the glycan structures they generate. Over the past decade, several components of protein glycosylation pathways have been investigated in pathogenic fungi, yet our understanding remains incomplete. Functional studies have been more extensive in human pathogens such as A. fumigatus, A. nidulans, C. albicans, and C. neoformans than in plant pathogens such as F. graminearum and M. oryzae (Fig. 1, Table 2). Notably, glycosylation research in human pathogens began earlier: in C. albicans, O- and N-glycogenes such as CaPMT1 and CaMNT1 were characterized in the late 1990s, followed by CaOCH1, CaMNN5, and ER α-glucosidases in the mid-2000s. By contrast, functional analyses in plant pathogenic fungi began nearly a decade later, with N-glycogenes in F. graminearum first reported in 2009, and subsequent work in F. oxysporum and M. oryzae emerging mainly after 2014. This temporal gap highlights the longer history of glycosylation research in human pathogens, and its recent expansion in plant pathogens.
Comparative analyses have revealed both conserved and divergent functional roles of glycogenes. PMTs are universally required for cell wall integrity, morphogenesis, and virulence, but their essentiality varies. C. albicans CaPMT2 and A. fumigatus PMT2 are indispensable for viability (Mouyna et al., 2010; Prill et al., 2005), whereas orthologs in B. cinerea or M. oryzae primarily affect adhesion and invasive growth (González et al., 2013, 2014; Guo et al., 2016). Similarly, OCH1 contributes to cell wall stability across fungi, yet its role in virulence differs markedly. In C. albicans, the CaOCH1 mutant lacking the α1,6-linked polymannose backbone was attenuated in virulence (Bates et al., 2006), whereas the OCH1 mutant in A. fumigatus showed no phenotypic differences from the wild type (Kotz et al., 2010). ALG genes, responsible for N-glycan precursor biosynthesis, also illustrate striking species-specific variations: complete loss of ALG2 in S. cerevisiae is lethal (Giaever et al., 2002), while deletion of MgALG2 in Mycosphaerella graminicola did not abolish viability but impaired cell wall integrity and morphogenesis (Motteram et al., 2011). Likewise, ER α-glucosidases (CWH41 and ROT2) in human pathogens mainly affect growth and cell wall composition (Mora-Montes et al., 2007; Mota et al., 2025; Zhang et al., 2008), whereas their counterparts in F. graminearum also regulate secondary metabolism and sexual reproduction (Moon et al., 2025). Taken together, these comparisons highlight that although glycogenes share core functions in protein maturation and cell wall integrity, their phenotypic outputs diverge across lineages, reflecting host-specific adaptations.
Despite relatively extensive genetic studies, direct evidence linking glycogenes to their corresponding glycan structures remains limited in pathogenic fungi. As summarized in Table 2, most reports focus on phenotypic outcomes rather than directly confirming glycan products, and only a subset employed structural analyses such as high-performance liquid chromatography (HPLC), matrix-assisted laser desorption/ionization time-of-flight mass spectrometry (MALDI-TOF MS), nuclear magnetic resonance (NMR) spectroscopy, or capillary electrophoresis. For example, in F. graminearum, Alg12 and Alg3 were confirmed to synthesize Dol-PP-GlcNAc2Man8 and Dol-PP-GlcNAc2Man6, respectively (Moon et al., 2025). In Fusarium oxysporum, Galf chain biosynthesis mediated by N-acetylglucosaminyltransferase 2 (Gnt2) was validated through size-exclusion chromatography combined with diffusion-ordered spectroscopy (DOSY) (Lopez-Fernandez et al., 2015). Similarly, in M. graminicola, Alg2 was implicated in (GlcNAc)2M3 synthesis based on analyses using MALDI-TOF MS (Motteram et al., 2011), while in A. fumigatus, structural studies confirmed that Och1 mediates outer chain mannose addition and Galf incorporation in GalfMan6GlcNAc2 biosynthesis (Kotz et al., 2010). Despite these advances, glycomic analyses remain scarce compared with gene deletion studies, leaving many predicted glycan structures unverified.
Importantly, fungal glycans often contain unique structural features absent from mammalian systems, such as polymannose outer chains in C. albicans (Bates et al., 2006), Galf residues in Aspergillus spp. (Henry et al., 2019; Kadooka et al., 2022; Onoue et al., 2018; Schmalhorst et al., 2008), or xylose-extended mannose branches in C. neoformans (Thak et al., 2020). These fungal-specific glycans are integral to cell wall integrity, morphogenesis, and immune evasion, thereby representing selective vulnerabilities. Because of their structural distinctiveness and absence in the host, fungal glycan epitopes constitute promising antifungal targets.
Glycoproteins in fungal virulence and development
Glycogenes are highly conserved across eukaryotes and are therefore less suitable as selective drug targets. In contrast, fungal glycoproteins—particularly those unique to fungal pathogens—often exhibit species-specific features that directly contribute to virulence, making them promising candidates for antifungal intervention. Similar to fungal-specific glycan structures, these glycoproteins operate at the host–pathogen interface, providing selective targets that minimize potential off-target effects in hosts.
Recent advances in glycoproteomics have enabled the large-scale identification of glycoproteins involved in cell wall integrity, signaling, and host–pathogen interactions. Most studies have focused on F. graminearum (Moon et al., 2025; Yu et al., 2016), F. oxysporum (Xu et al., 2020b), Colletotrichum graminicola (Mei et al., 2023), M. oryzae (Chen et al., 2020), B. cinerea (González et al., 2014), A. nidulans (Rubio et al., 2016), Aspergillus niger (Wang et al., 2012), and C. albicans (Léger et al., 2016). Across these diverse fungi, glycoproteomics consistently reveal common features that are critical for both fungal development and pathogenicity. Most glycoproteins are localized to the cell wall or secretory pathway, underscoring their roles at the host–pathogen interface. They typically function as adhesins, hydrolytic enzymes, or glycosylphosphatidylinositol (GPI)-anchored mannoproteins, thereby promoting adhesion, cell wall remodeling, and immune evasion (Léger et al., 2016; Rubio et al., 2016; Wang et al., 2012). Comparative analyses also demonstrate that glycoproteins rely heavily on N- and O-linked glycosylation for proper folding and stability via ER quality control (ERQC). Disruption of these processes often results in protein misfolding, defective secretion, and attenuated virulence (Chen et al., 2020; González et al., 2014; Mei et al., 2023; Moon et al., 2025). Importantly, glycoproteomic profiling has highlighted both conserved and species-specific repertoires, reflecting shared as well as specialized pathogenic strategies.
Building on these large-scale approaches, functional characterization of individual glycoproteins has further clarified their biological significance. As summarized in Table 3, these studies reveal the multifaceted roles of glycoproteins across diverse pathogenic fungi. Adhesins, such as Hwp1 and the Als family in C. albicans (Hoyer et al., 2008; Kamai et al., 2002; Sharkey et al., 1999; Zhao et al., 2004, 2005), and lysine motif (LysM) domain–containing proteins, including Slp1 in M. oryzae (Chen et al., 2014; Jones et al., 2021) and LysM (Crumière et al., 2025) in B. cinerea, are directly involved in host adhesion, immune evasion, and infection initiation. Secreted enzymes, such as endopolygalacturonases (Pg1 in B. cinerea and Pcipg2 in Phytophthora capsici) (González et al., 2014; ten Have et al., 1998; Sun et al., 2009), act as virulence factors by degrading host cell walls. Numerous glycoproteins also contribute to fundamental cellular processes, including vegetative growth, cell wall integrity, sporulation, and sexual reproduction, exemplified by Ecm33, Cwm1, and chitin synthases in F. graminearum (Fernando et al., 2019; Kim et al., 2009; Zhang et al., 2019a). ER chaperone (e.g., Cnx1) and glucosidase (e.g., Gtb1) enzymes associated with glycosylation or ERQC (Chen et al., 2020) further highlight the reliance of fungal development and pathogenicity on proper folding and trafficking. Notably, while some glycoproteins are essential for viability (e.g., phosphatases, Hsp70 family proteins) (Liu et al., 2017; Yu et al., 2014; Yun et al., 2015), others display defects only during host infection, underscoring their specialized roles in pathogenesis (Cadieux et al., 2013; Nigam et al., 2003; Pietrella et al., 2002; Son et al., 2011; Yin et al., 2018a; Yun et al., 2015). Together, these studies illustrate how glycoproteins function at the interface of fungal physiology and pathogenicity, reinforcing their significance as selective and promising targets for antifungal intervention.
While a comprehensive, network-level framework linking the hierarchical layers of glycosylation has yet to be fully established, emerging evidence suggests functional interconnections across these levels. Here, we outline glycoprotein-centered virulence networks (Fig. 2), proposing that glycosylation governs fungal pathogenicity through a multilayered system integrating ERQC, secretory trafficking, cell wall biogenesis, and the activity of surface and secreted glycoproteins at the host interface. To illustrate these concepts, we highlight two representative contexts: plant pathogenic fungi and human pathogenic fungi, focusing on C. neoformans and C. albicans.
Plant pathogenic fungi
In plant pathogens, glycosylation underpins penetration and invasive growth by stabilizing and localizing key glycoproteins required for appressorium formation, cell wall remodeling, immune evasion, and oxidative stress tolerance. Functionally, these glycoproteins can be broadly grouped into five categories: ERQC, autophagy, cell wall integrity, reactive oxygen species (ROS) resistance, and defense against host-derived chitinases and other antifungal enzymes (Fig. 2A).
The ERQC system is indispensable for proper protein folding and stability, thereby shaping fungal virulence. In M. oryzae, Cnx1 and Gtb1 are required for vegetative growth, conidiation, and invasive hyphal growth (Chen et al., 2020). In C. graminicola, the ERQC components such as Cnx1 likewise ensure effector stability and secretion, underscoring their central role in host–pathogen interactions (Mei et al., 2023).
Vesicle trafficking and autophagy are tightly interconnected, coordinating fungal development, stress adaptation, and secondary metabolism. In F. graminearum, the SNARE protein Syn2 is required for sexual development (Hong et al., 2010), while the vacuolar sorting factor Vps74 contributes to vegetative growth, conidiation, sexual reproduction, and mycotoxin production (Kim et al., 2015b). Autophagy-related factors intersect with pathogenic processes: Atg27 mediates selective autophagy pathways such as mitophagy, pexophagy, and the cytoplasm-to-vacuole targeting (Cvt) pathway, influencing development and mycotoxin production (Lv et al., 2017). Similarly, the subtilisin-like protease Prb1 contributes to mycotoxin biosynthesis, lipid metabolism, and environmental stress responses, further linking autophagy to virulence (Xu et al., 2020a).
Cell wall integrity is fundamental for fungal survival and virulence, serving as a dynamic barrier that both protects against stresses and mediates host–pathogen interactions. In F. graminearum, chitin synthases Chs7 and Chs5 are required for growth, septum formation, and perithecium development (Kim et al., 2009). Arb1, an ATP-binding protein, plays a broader role by contributing to infective growth, oxidative stress resistance, cell wall maintenance, mycotoxin production, and sexual/asexual development (Yin et al., 2018a). Cwm1, a cell wall mannoprotein, is essential for vegetative growth and the cell wall integrity (Zhang et al., 2019a). In M. oryzae (Liu et al., 2020), a GPI-anchored protein Gpi7 drives penetration, invasive growth, and cell wall biogenesis.
Secretory and cell wall-associated proteins act directly at the pathogen-host interface. In F. graminearum, KatG2, a wall-localized peroxidase, detoxifies ROS during invasion (Guo et al., 2019), while Glx, a plasma-membrane glyoxal oxidase, contributes to both ROS detoxification and to mycotoxin production (Song et al., 2016). Ecm33, a GPI-anchored protein, is required for maintaining proper cell wall morphology and integrity (Fernando et al., 2019). Multiple LysM effectors act as secretory proteins that counteract host defense enzymes: Lysm1 in B. cinerea, Slp1 in M. oryzae, and Scp1 in Lasiodiplodia theobromae (Chen et al., 2014; Crumière et al., 2025; Jones et al., 2021; Peng et al., 2022). Notably, Lysm1 in B. cinerea also scavenges ROS, underscoring its dual protective role.
Human pathogenic fungi
In C. neoformans, cell surface glycoproteins such as Cig1, Mp65, and Mp84 (Fig. 2B) act as key antigens that facilitate adhesion, modulate immune responses, and promote persistence (Cadieux et al., 2013; Pietrella et al., 2002; Teixeira et al., 2014). Their antigenic properties enhance host recognition while simultaneously enabling immune evasion. Capsule formation is also a major virulence determinant: the polysaccharide capsule, composed primarily of glucuronoxylomannan (GXM)- and galactoxylomannan (GalXM)-rich capsule blocks immune recognition, inhibits phagocytosis, and modulates cytokine signaling. Cmp1 regulates capsule biosynthesis (Han et al., 2020), while Plb1 facilitates intracellular infection and alters fungal cell morphology during infection (Evans et al., 2015; Thak et al., 2020). Taken together, these glycoproteins and capsule-associated factors orchestrate the multifaceted virulence program of C. neoformans.
In C. albicans, several glycoproteins play pivotal roles in virulence by orchestrating adhesion, morphogenesis, and hyphal development (Fig. 2C). MP65 mediates fungal cell adhesion and hyphal morphogenesis, thereby promoting initial host colonization and the morphological transitions essential for invasive growth (Sandini et al., 2007). Hwp1, a hypha-specific glycoprotein, supports hyphal development and acts as a substrate for host transglutaminases, enhancing stable adhesion to epithelial surfaces (Sharkey et al., 1999; Staab et al., 2013). The Als family proteins, particularly Als1, Als2, and Als3, are major cell surface adhesins that mediate binding to host tissues and abiotic substrates, facilitating biofilm formation and immune evasion (Hoyer et al., 2008; Kamai et al., 2002; Zhao et al., 2004). Collectively, these glycoproteins underscore the multifactorial nature of C. albicans pathogenicity, linking surface adhesion and morphogenetic plasticity to host invasion and persistence.
Over the past decade, numerous studies have highlighted the fundamental roles of glycosylation in fungal infection. It is now evident that fungal glycosylation is intricately involved in diverse aspects of pathogenicity, including growth, morphogenesis, cell wall biogenesis, protein secretion, and host immune modulation. Future research should therefore move beyond gene-centric approaches toward a more integrated understanding of how glycosylation drives fungal virulence. Recent advances integrating proteomics, transcriptomics, and glycomics in species such as F. graminearum, M. oryzae, C. graminicola, and C. neoformans have begun to unravel the multilayered roles of glycosylation in fungal virulence (Chen et al., 2020; Mei et al., 2023; Moon et al., 2025; Mota et al., 2025). Nevertheless, an overarching framework that links the hierarchical layers of glycosylation with fungal development and pathogenesis has yet to be established. In this review, we summarize the current knowledge and emphasize the need for such integrative models.
Despite these advances, significant challenges remain. The first major challenge is the comprehensive profiling of glycan structures in pathogenic fungi. While analogous efforts in bacteria, viruses, and cancer research have progressed rapidly, fungal systems remain relatively underexplored. For instance, in SARS-CoV-2, the high-mannose and complex glycan shield of the spike protein has been targeted using glycan-binding lectins and antibodies (Klevanski et al., 2024). In bacteria, the O-antigen glycans of Gram-negative species have emerged as promising vaccine and antibody targets (Zhang et al., 2009). In cancer, tumor-associated carbohydrate antigens (TACAs) such as sialylated epitopes and aberrant N-glycan branching are under active investigation as therapeutic targets (Rashidijahanabad and Huang, 2020; Zhou et al., 2025). In contrast, antifungal strategies aimed at glycan structures remain at an early stage. Although resources such as the GlycoShape database provide approximations of glycoprotein structures (Ngolong Ngea et al., 2021), the development of clinically relevant antifungal targets will require precise and systematic profiling of pathogen-specific glycans. Recent studies have begun to characterize glycan landscapes in C. neoformans, F. graminearum, and A. fumigatus (Moon et al., 2025; Onoue et al., 2018; Park et al., 2012). Moving forward, these efforts must expand to include comparative profiling of host glycomes in plants and animals. Such integrative analyses will help identify pathogen-specific glycan motifs, thereby providing a foundation for the rational design of antifungal agents.
The second challenge is to elucidate whether and how distinct glycan structures directly influence the functions of their cognate glycoproteins. Large-scale glycoproteomic studies have identified numerous glycoproteins associated with virulence (Chen et al., 2020; Mei et al., 2023; Moon et al., 2025; Yu et al., 2016), but a critical next step is to determine whether their functions depend simply on the presence of glycans, or on their specific structural features. The absence of glycosylation at a given site can be examined through site-directed mutagenesis of the corresponding amino acid residues (Guo et al., 2019; Mei et al., 2023; Peng et al., 2022). In contrast, assessing the contribution of glycan structure requires more advanced approaches, including LC–MS/MS of released glycans or glycopeptides, exoglycosidase digestion, and complementary techniques such as NMR or lectin-based profiling. Furthermore, experimental systems such as glycoengineering of fungal pathways, application of glycosidase inhibitors, and chemoenzymatic remodeling of glycoproteins can generate distinct glycoforms of the same protein. By comparing the activities, stability, and virulence-related properties of these glycoforms, researchers can directly evaluate the functional relevance of glycan structural variation. Such analyses will be essential for establishing whether particular glycan motifs represent viable and selective targets for antifungal drug development.
The online version contains supplementary material available at https://doi.org/10.71150/jm.2510003.
Fig. 1.
Summary of the pathogenic contributions of glycogenes associated with N- and O-glycosylation across representative fungal species. (A) Pathogenic contribution map of proteins involved in (A) N-glycosylation and (B) O-glycosylation pathways across fungal species. Species are color-coded; filled circles indicate genes with demonstrated pathogenic relevance, whereas open circles represent genes studied without pathogenic association. Abbreviations: Fg, Fusarium graminearum; Mo, Magnaporthe oryzae; Afu, Aspergillus fumigatus; Ca, Candida albicans; Cn, Cryptococcus neoformans. Filled circles indicate glycogenes that contribute to virulence, whereas unfilled circles represent glycogenes that have been characterized but do not affect virulence. Each positions indicate pathway step, not physical interaction.
jm-2510003f1.jpg
Fig. 2.
Potential regulatory mechanisms of glycoproteins in fungal virulence. Functional analyses of glycogenes have been conducted in both human and plant pathogenic fungi. Conserved glycoproteins contribute to key processes such as protein maturation, cell wall integrity, morphogenesis, and virulence, yet their functional impacts vary in a species-specific manner across fungal lineages. (A) In plant pathogenic fungi, glycosylation regulates penetration and invasive growth by stabilizing and positioning key glycoproteins required for invasive growth, cell wall remodeling, immune evasion, and oxidative stress tolerance. These proteins can be grouped into ER quality control (ERQC; Cnx1, Gtb1, and Ecm33), autophagy and vesicle trafficking (Atg27, Syn2, Vps74, and Prb1), and cell wall integrity factors (Cwm1, Chs7, Chs5, and Arb1), all contributing to fungal development, sporulation, and mycotoxin production. Additional cell wall–localized or secretory proteins, such as Gpi7, KatG2, Glx, and LysM effectors (LysM1, Slp1, Scp1), contribute to invasive growth by scavenging host-derived reactive oxygen species and protecting fungal cell walls from plant chitinases. (B) In the human pathogen C. neoformans, several glycoproteins function as antigens and virulence determinants at the cell surface. Cig1, Mp65, and Mp84 act as immunogenic proteins that mediate host–pathogen interactions, while Cmp1 regulates capsule formation and Plb1 contributes to intracellular infection and morphological adaptation. Capsule polysaccharides further support immune evasion and pathogen survival. (C) In C. albicans, glycoproteins orchestrate adhesion, morphogenesis, and hyphal development. MP65 promotes fungal adhesion and morphogenetic transitions; Hwp1 facilitates hyphal development and stable adhesion to host tissues; and the Als family proteins (Als1, Als2, and Als3) function as major surface adhesins driving biofilm formation and immune evasion. This figure was created using BioRender.
jm-2510003f2.jpg
Table 1.
TFs regulating glycogenes
Species Gene Type Description Regulated genes Direct binding Reference
C. albicans CaACE2 Zinc-finger RAM (regulation of ACE2 TF and polarized morphogenesis) signaling network PMTs Not studied Cantero and Ernst (2011); van Wijlick et al. (2016)
B. cinerea BcFTG1 Zinc-finger fungal TF containing the GAL4 domain ALG8 Not studied Yang et al. (2024)
H. polymorpha HpHAC1 bZIP Major unfolded protein response (UPR) ALG5, MNN2, MNN4, KTR1, OST1,OST2, OST4, OCR5, SWP1, WBP1 Not studied Moon et al. (2015)
A. niger hacA bZIP Major unfolded protein response (UPR) ALG6, ALG8, ALG2, ALG3, DPM1,DPM2, ALG5, ALG7, RFT1, ALG9, SEC59, ALG12, ALG10, OST1, WBP1,OST2, OST3, SST3 Not studied Carvalho et al. (2012)
C. neoformans HXL1 bZIP HAC1, XBP1-like gene 1 UPR genes (WBP1, PMT1, PMT2, PMT4, WBP1, ALG7, OST1) Not studied Cheon et al. (2011); Glazier et al. (2015)
Mediate the unfolded protein response (UPR) pathway
M. robertsii MrpacC C2H2-type zinc-finger pH-responsive TF STT3, OST3, PMT4 STT3, OST3, PMT4 Huang et al. (2015)
M. acridum MaSte12 C2H2-type zinc-finger Direct target and functions downstream of the mitogen-activated protein kinase (MAPK) Fus3/Kss1 OCH1 Not studied Wei et al. (2017)
Table 2.
N&O-Glycogenes and glycan structures discussed in this review
Species Gene Virulence (O = affected; X = no effect) Biological function(s) Glycan structure Reference
B. cinerea bcpmt1 O Morphogenesis Not studied González et al. (2013, 2014)
Fungal adherence
Cell wall integrity
bcpmt2 O Morphogenesis Not studied González et al. (2013, 2014)
Fungal adherence
Cell wall integrity
bcpmt4 O Morphogenesis Not studied González et al. (2013, 2014); Plaza et al. (2025)
Fungal adherence
Cell wall integrity
C. graminicola Cgalg3 O ER homeostasis Not studied Mei et al. (2023)
Fungal growth
F. graminearum RFT1 O Mycotoxin synthesis Not studied Moon et al. (2025)
Vegetative growth
Conidiation
Sexual development
CWH41 (MOGS) X Mycotoxin synthesis Not studied
Conidiation
Sexual development
ROT2 (GLS2, GANAB) X 6 orthologs showed different phenotypes Not studied
Mycotoxin synthesis
Vegetative growth
Conidiation
Sexual development
ALG5 O Vegetative growth Not studied
Sexual development
ALG6 O Vegetative growth Not studied
Sexual development
ALG8 O Mycotoxin synthesis Not studied
Vegetative growth
Sexual development
PSA1 (MPG) O Mycotoxin synthesis Not studied
Vegetative growth
Conidiation
Sexual development
MNS1 (MAN1B) O 2 orthologs showed different phenotypes Not studied
Mycotoxin synthesis
Conidiation
MNL2 (MAN1A) O 4 orthologs showed different phenotypes Not studied
Mycotoxin synthesis
Vegetative growth
Conidiation
PMT1, 5, 7 O Vegetative growth Not studied
Conidiation
Sexual development
MNN14 X Mycotoxin synthesis Not studied
Vegetative growth
Conidiation
ALG1 X Vegetative growth Not studied
ALG11 O Mycotoxin synthesis Not studied
Vegetative growth
Conidiation
Sexual development
ALG12 O Vegetative growth Biosynthesis of the Dol-PP-GlcNAc2Man8 (HPLC)
Sexual development
ALG2 o Mycotoxin synthesis Not studied
Vegetative growth
Conidiation
Sexual development
ALG3 O Mycotoxin synthesis Biosynthesis of the Dol-PP-GlcNAc2Man6 (HPLC)
Vegetative growth
Conidiation
Sexual development
ALG9 X Mycotoxin synthesis Not studied
Vegetative growth
HOC1 O Mycotoxin synthesis Not studied
OCH1 O Mycotoxin synthesis Not studied
Vegetative growth
Conidiation
KTR5, KTR7 O Mycotoxin synthesis Not studied
Vegetative growth
Conidiation
Sexual development
KTR4 O Vegetative growth Not studied
Conidiation
KRE2, KTR1, 2, 3, 6 X Mycotoxin synthesis Not studied
Vegetative growth
Sexual development
MNT4 X Mycotoxin synthesis Not studied
Conidiation
MNN1, MNT3 X Mycotoxin synthesis Not studied
MNN10 X Conidiation Not studied
Sexual development
MNN11 X Mycotoxin synthesis Not studied
Vegetative growth
Conidiation
Sexual development
MNN2, 5 X Conidiation Not studied
MNN4 O Mycotoxin synthesis Not studied
Vegetative growth
VAN1 X Mycotoxin synthesis Not studied
Vegetative growth
Sexual development
ANP1 O Mycotoxin synthesis Not studied
Vegetative growth
Conidiation
Sexual development
MNN9 O Vegetative growth Not studied
Sexual development
STT3 O Not changed Not studied
SWP1 X Conidiation Not studied
WBP1 O Mycotoxin synthesis Not studied
Vegetative growth
Conidiation
F. oxysporum OCH1 O Fungal growth Not studied Li et al. (2014)
Cell wall integrity
hyphal adhesion
GNT2 O Conidium size and morphology Biosynthesis of the Galactofuranose chains Lopez-Fernandez et al. (2015)
Oxidative stress resistance (size-exclusion chromatography with Diffusion Ordered Spectroscopy (DOSY))
cell wall organization, biogenesis and remodelling hyphal fusion
Secretion of trafficking vesicles
F. oxysporum f. sp. cucumerinum PMT1 O Fungal growth Not studied Li et al. (2014)
Cell wall integrity
Conidiation
PMT2 O Fungal growth Not studied Li et al. (2014)
Cell wall integrity
Conidiation
PMT4 O Fungal growth Not studied Xu et al. (2020b)
Cell wall integrity
Conidiation
M. graminicola MgAlg2 O Cell wall integrity Biosynthesis of the (GlcNAc)2M3 Motteram et al. (2011)
Yeast-like to filamentous growth switch (MALDI-ToF-MS)
M. oryzae GLS1 O Mycelial growth Not studied Chen et al. (2020)
Conidiation
Invasive hyphal growth
GLS2 O Mycelial growth Not studied Chen et al. (2020)
Conidiation
Invasive hyphal growth
GTB1 O Mycelial growth Not studied Chen et al. (2020)
Conidiation
Invasive hyphal growth
ALG3 O Mycelial growth Not studied Chen et al. (2014)
Conidiation
Invasive growth
Oxidative stress resistance
PMT2 O Fungal adhesion Not studied Guo et al. (2016)
Conidial germination
cell wall integrity
Invasive hyphae growth
PMT4 O Hyphal growth Not studied Pan et al. (2019)
Conidiation
Penetration and biotrophic invasion
ALG9 O Conidial production Not studied Zhang et al. (2025)
Appressorium formation
Responses to stressors
P. capsici PcSTT3B O Vegetative growth Not studied Cui et al. (2023)
Sporangial release rate
Zoospore production
P. digitatum pdpmt2 O Cell wall integrity Not studied Harries et al. (2015)
Conidiogenesis
Sensitivity to fungicide
U. maydis gls1 O Initial stages of biotrophic growth Not studied Fernández-Álvarez et al. (2013)
gas1 O Intracellular expending Not studied Schirawski et al. (2005)
gas2 O Intracellular expending Not studied Fernández-Álvarez et al. (2013)
pmt4 O Appressorium formation Not studied Fernández-Álvarez et al. (2009)
Plant cuticle penetration
V. dahliae VdOCH1 O Hyphal growth Not studied Zhang et al. (2019b)
Conidia production
Microsclerotia formation
Cell wall integrity
VdSTT3 O Fungal development Not studied Su et al. (2018)
Hyphal growth
Glycoprotein secretion
A. fumigatus CWH41 X Cell wall integrity (CWI) Not studied Zhang et al. (2008)
Polarity
Septation
Conidiation
AMS1 Not studied Conidial formation Not studied Li et al. (2009)
Polarity
Septation
OCH1 O Sporulation Biosynthesis of the Outer glycan chain Kotz et al. (2010)
(Capillary electrophoresis DNA Sequencer)
PMT1 X No discernible phenotype Not studied Mouyna et al. (2010)
ΔPMT1/PMT4 was lethal
PMT2 O Essential Not studied Mouyna et al. (2010)
PMT4 X Mycelial growth Not studied Mouyna et al. (2010)
Conidiation
Sensitivity to echinocandins
ΔPMT1/PMT4 was lethal
ANPA X Mycelial growth Core-mannan structure of fungal-type galactomannan (FTGM) Kadooka et al. (2022)
Conidia formation (1H-NMR spectroscopy)
MNN9 X No discernible phenotype Not involved in the synthesis of the FTGM α-core-mannan Du et al. (2019); Kadooka et al. (2022)
Apical growth and polarity (1H-NMR spectroscopy)
No significant role in the biosynthesis of N-glycans
(MALDI-TOF MS)
VAN1 X No discernible phenotype Not involved in the synthesis of the FTGM α-core-mannan Kadooka et al. (2022)
(1H-NMR spectroscopy)
KTR4 O Polarized growth Polymerization galactomannan Henry et al. (2019)
Conidiation (HPLC analysis)
Conidial viability
KTR7 O Polarized growth Polymerization galactomannan Henry et al. (2019)
Conidiation (HPLC analysis)
Conidial viability
CMSA Not studied Mycelial growth Biosynthesis of the FTGM core-mannan structure Onoue et al. (2018)
sensitive to antifungal agents (Proton nuclear magnetic resonance (1H-NMR) spectroscopy)
CMSB Not studied Mycelial growth Biosynthesis of the FTGM core-mannan structure Onoue et al. (2018)
sensitive to antifungal agents (Proton nuclear magnetic resonance (1H-NMR) spectroscopy)
MNT1 O Cell wall stability Not studied Wagener et al. (2008)
MNN2 Not studied No discernible phenotype ΔMNN2/MNN5 Kadooka et al. (2024)
ΔMNN2/MNN5 exhibited a growth defect and abnormal conidial formations Biosynthesis of the outer chain like α-(1→6)-linked mannan
(1H-NMR analysis)
MNN5 Not studied No discernible phenotype ΔMNN2/MNN5 Kadooka et al. (2024)
ΔMNN2/MNN5 exhibited a growth defect and abnormal conidial formations Biosynthesis of the outer chain like α-(1→6)-linked mannan
(1H-NMR analysis)
STT3 Not studied Fungal growth Not studied Li et al. (2011)
Cell wall integrity (CWI)
GALF O Morphology and growth Biosynthesis of the GalfMan6GlcNAc2 Schmalhorst et al. (2008)
Susceptible to drugs (Four-capillary electrophoresis DNA sequencer)
A. nidulans ALG7 Not studied Failed to delete Not studied Gerhardt et al. (2025)
ALG6 (ALGF) Not studied Not studied Not studied Anyaogu et al. (2021); Gerhardt et al. (2025)
AMS1 X No visible effect on growth or morphology Not studied Anyaogu et al. (2021); Eades et al. (1998)
Recycling macromolecular components
AN5748 Not studied Fungal growth Not studied Gerhardt et al. (2025)
Sporulation
Resistance to sorbitol and tunicamycin
PMTA Not studied Fungal growth Not studied Oka et al. (2004)
Cell wall formation
ΔPMTA/PMTC lethal
PMTB Not studied Conidiation Not studied Goto et al. (2009)
Polarity maintenance
ΔPMTB/PMTC lethal
PMTC Not studied Fungal growth Not studied Goto et al. (2009)
Cell wall integrity
Osmotic stabilization
ΔPMTA/PMTC, ΔPMTB/PMTC lethal
ALG1 Not studied Failed to delete Not studied Gerhardt et al. (2025)
ALG2 Not studied Fungal growth Not studied Gerhardt et al. (2025)
Sporulation
ALG11 Not studied Failed to delete Not studied Gerhardt et al. (2025)
ALG3 (ALGC) Not studied Sensitive to tunicamycin and calcium stress Biosynthesis of the Man5-7GlcNAc2 Anyaogu et al. (2021); Gerhardt et al. (2025)
(UHPLC-FLR-MS)
ALG9 (ALGI) Not studied Sensitive to tunicamycin and calcium stress Not studied Anyaogu et al. (2021); Gerhardt et al. (2025)
ALG12 (ALGL) Not studied Failed to delete Not studied Anyaogu et al. (2021); Gerhardt et al. (2025)
ALG13 Not studied Failed to delete Not studied Gerhardt et al. (2025)
C. albicans CaCWH41 O Growth rates Not studied Mora-Montes et al. (2007)
Cell wall composition
CaROT2 o Growth rates Not studied Mora-Montes et al. (2007)
Cell wall composition
CaMNS1 O Growth rates Not studied Mora-Montes et al. (2007)
Cell wall composition
CaPMT2 O Essential Not studied Prill et al. (2005)
CaPMT4 O Fungal growth Not studied Prill et al. (2005)
Morphogenesis
Antifungal resistance.
CaPMT5 X Fungal growth Not studied Prill et al. (2005)
Morphogenesis
Antifungal resistance.
CaPMT1 O (heterozygous)/ Hyphal morphogenesis Not studied Eades et al. (1998); Timpel et al. (2000)
X (homozygous) Supersensitivity to the antifungal agents
CaPMT6 O Morphogenesis Not studied Timpel et al. (2000)
Antifungal sensitivities
CaOCH1 O Temperature-sensitive growth Biosynthesis of the N-glycan outer chain Bates et al. (2006)
Cellular aggregation (ES-MS analysis, Gas chromatography-mass spectrometry)
Host-fungal interaction
CaMNT1 O Adhesion Biosynthesis of the second mannose to O-glycan Buurman et al. (1998)
(Biogel-P4 chromatography)
CaMNN5 O Hyphal growth Biosynthesis of the N-linked mannan branches Bai et al. (2006)
Iron homeostasis (Thin-layer chromatography and TLC plate to autoradiography)
Cell wall integrity
Morphogenesis
CaMNT4 Not studied No discernible phenotype Not studied Mora-Montes et al. (2010)
ΔMNT3/MNT4/MNT5 exhibited severe growth defect
ΔMNT4/MNT5, ΔMNT3/MNT4/MNT5 altered cell wall composition
CaMNT5 O (ΔMNT3/MNT5) No discernible phenotype Not studied Mora-Montes et al. (2010)
ΔMNT3/MNT4/MNT5 exhibited severe growth defect
ΔMNT4/MNT5, ΔMNT3/MNT4/MNT5 altered cell wall composition
CaMNT3 O (ΔMNT3/MNT5) No discernible phenotype Not studied Mora-Montes et al. (2010)
ΔMNT3/MNT5, ΔMNT3/MNT4/MNT5 exhibited severe growth defect
ΔMNT3/MNT5, ΔMNT4/MNT5, ΔMNT3/MNT4/MNT5 altered cell wall composition
C. neoformans MNS1 (ΔMNS1/MNS101) Sensitive to ER stress (tunicamycin, DTT) Remove mannose from GlcNAc2Man9 to GlcNAc2Man8 Mota et al. (2025)
Sensitive to ER stress cell wall integrity stress (HPLC and MALDI-TOF)
ΔMNS1/MNS101 growth defect
MNS101 (ΔMNS1/MNS101) Sensitive to ER stress (tunicamycin, DTT) Remove mannose from (> M10) glycan or M8 further trimming Mota et al. (2025)
Sensitive to ER stress cell wall integrity stress (HPLC and MALDI-TOF)
ΔMNS1/MNS101 growth defect
MNL1 X No discernible phenotype Involved in targeting misfolded proteins for ERAD Mota et al. (2025)
rather than in normal N-glycan processing
(HPLC, MALDI-TOF)
MNL2 X No discernible phenotype Involved in targeting misfolded proteins for ERAD Mota et al. (2025)
rather than in normal N-glycan processing
(HPLC, MALDI-TOF)
PMT4 O Fungal growth Not studied Olson et al. (2007); Willger et al. (2009)
Cell wall integrity
Cell morphology
PMT2 O Essential Not studied Willger et al. (2009)
PMT1 O Cell morphology and integrity Not studied Willger et al. (2009)
ΔPMT1/PMT4 is synthetically lethal,
ALG3 O Macrophage cell death Biosynthesis of the Dol-PP-GlcNAc2Man6 Thak et al. (2020)
(HPLC, MALDI-TOF)
OCH1 Slightly attenuated, No discernible phenotype Addition of a single α1,6-linked mannose residue to the Man8GlcNAc2 core Park et al. (2012)
not critical (HPLC, MALDI-TOF)
ALG9 O Sensitive to SDS, fluconazole Biosynthesis of the Dol-PP-GlcNAc2Man7 Thak et al. (2020)
(HPLC, MALDI-TOF)
MNN2 Not studied No discernible phenotype Biosynthesis of the elongated outer chain N-glycan Park et al. (2012)
(HPLC, MALDI-TOF)
KTR3 Not studied Cell wall stability Not involved in the processing of N-glycans (M6–10, X1M6–10) Park et al. (2012)
(HPLC, MALDI-TOF)

Man9GlcNAc2 : Glycan compositions with 9 mannose and two N-acetylglucosamine residues.

X1M6–10: Extended high-mannose glycans containing one Xylose and 6–10 mannose residues.

FTGM: Fungal-type galactomannan.

Table 3.
Glycoproteins in fungal virulence and development
Species Gene Description Phenotypic defect(s) Reference
B. cinerea bcsun1 Member of the β-glucosidase SUN family Fungal morphogenesis Pérez-Hernández et al. (2017)
(secreted glycoprotein) Cell Wall Integrity
Reproductive structures formation
ROS sensitivity
bcpg1 Endopolygalacturonase Studied only for virulence González et al. (2014); ten Have et al. (1998)
bclysm1 Lysin motif domains Protects hyphae against degradation by external chitinases Crumière et al. (2025)
Prevent them from inducing ROS
Infection initiation
Adhesion to host
C. graminicola Cgcnx1 ER chaperone (Site directed mutagenesis) Mei et al. (2023)
Vegetative growth
Effector stability and secretion
F. graminearum CFEMN1 Contain a CFEM domain Interact with extracellular binding Zuo et al. (2022)
proteins from host
TUB2 β2-tubulin Vegetative growth Liu et al. (2013);
Sensitive to carbendazim Zhao et al. (2014)
ADE5 Phosphoribosylamine-glycine ligase Adenine auxotrophy Kim et al. (2007)
Vegetative growth
Sexual reproduction
CHS7 Chitin synthase Vegetative growth Kim et al. (2009)
Septa formation
Perithecia formation
CHS5 Chitin synthase Vegetative growth Kim et al. (2009)
Septa formation
Perithecia formation
VPS74 Vacuolar protein sorting Vegetative growth Kim et al. (2015b)
Conidiation
Sexual development
Mycotoxin production
SYN2 Syntaxin-like SNARE genes Self and female fertility Hong et al. (2010)
Localized in plasma membranes and septa
ERG9 Ergosterol biosynthesis Essential Yun et al. (2014)
PRB1 Subtilisin-like protease belonging to proteinase K-like subfamily Mycotoxin production Xu et al. (2020a)
Responses to environmental stimuli
Lipid metabolism
Autophagy regulation
ATG27 Autophagy-related genes Vegetative growth Lv et al. (2017)
Sporulation
Mycotoxin production
Autophagy regulation
ILV6 Acetohydroxyacid synthase BCAA-auxotroph Liu et al. (2015)
Vegetative growth
Pigmentation
Mycotoxin production
ILV2 Acetohydroxyacid synthase BCAA-auxotroph Liu et al. (2015)
Vegetative growth
Conidial formation
Pigmentation
Mycotoxin production
GPB1 G protein Mycotoxin production Yu et al. (2008)
Chitin accumulation
RAB11 Ras-related proteins in brain Polarized growth and/or exocytosis Zheng et al. (2015)
KAR Hsp70s Essential Liu et al. (2017)
SSC Hsp70s Essential Liu et al. (2017)
SSB Hsp70s Proper folding of β2-tubulin Liu et al. (2017)
Vegetative growth
Low temperature response
Vacuole fusion and endocytosis
Mycotoxin production
SSA Hsp70s Essential Liu et al. (2017)
CAP1 Cyclase-associated protein Vegetative growth Yin et al. (2018b)
Conidiogenesis
Mycotoxin production
KATG2 /FCA7 peroxidase (Site directed mutagenesis) Guo et al. (2019);
Scavenges ROS Lee et al. (2018)
Located on the cell wall of invading hyphal cells
ECM33 GPI-anchored protein Cell wall morphology Fernando et al. (2019)
PP2A Phosphatase Essential Yu et al. (2014)
STE7 kinase Vegetative growth Gu et al. (2015)
Conidiation
Mycotoxin production
Penetration
PHO8 Phosphatase Essential Yun et al. (2015)
CNA1 Phosphatase Essential Yun et al. (2015)
Fg07304 Phosphatase Essential Yun et al. (2015)
GIC1 Phosphatase Conidiation Yun et al. (2015)
PSR2 Phosphatase Vegetative growth Yun et al. (2015)
Pigmentation
Conidia morphology
ARC15 Phosphatase Only defects in virulence Yun et al. (2015)
GLX glyoxal oxidase Mycotoxin production Song et al. (2016)
localized to cell membrane
ARB1 ATP-binding protein Infective growth Yin et al. (2018a)
Oxidative stresses
Cell wall integrity
Mycotoxin production
Sexual and asexual development
SKP1 E3 ubiquitin ligases Essential Chen et al. (2023)
LIP50, TAG Lipases Only defects in virulence Kim et al. (2023); Zhang et al. (2016)
PCT1 Cytidylyltransferases Essential Wang et al. (2019)
FGSG_10825 Mating-type Sexual development Kim et al. (2015a)
OB031, HEX1 TF Only defects in virulence Son et al. (2011)
CMK1 Kinase Ascospore formation Wang et al. (2011)
ALDH (FGSG_02273) Aldehyde Dehydrogenase Essential Tang et al. (2023)
ALDH (FGSG_04194) Aldehyde Dehydrogenase Regulating acetaldehyde dehydrogenase activity Tang et al. (2023)
Accumulation of lipid droplets
Conidial germination
CWM1 Cell wall mannoprotein Vegetative growth Zhang et al. (2019a)
Cell wall integrity
L. theobromae LtScp1 Effectors, a LysM-containing protein (Site directed mutagenesis) Peng et al. (2022)
Interacted with chitinase VvChi4
M. oryzae CNX1 Calnexin Mycelial growth Chen et al. (2020)
Conidiation
Invasive hyphal growth
GTB1 Glucosidase II β-subunit Gas2 Mycelial growth Chen et al. (2020)
Conidiation
Invasive hyphal growth
SLP1 Secreted LysM domain protein (Site directed mutagenesis) Chen et al. (2014); Jones et al. (2021)
Sequesters chitin oligosaccharides
Apoplastic effectors
BAS4 Biotrophy-associated secreted protein 4 Apoplastic effectors Chen et al. (2014); Jones et al. (2021)
GPI7 Glycosylphosphatidylinositol Penetration and invasive growth Liu et al. (2020)
Cell wall biogenesis
P. capsici Pcipg2 Polygalacturonase Studied only for virulence Sun et al. (2009)
P. sojae GPI16 GPI transamidase component protein (Site directed mutagenesis) Zhang et al. (2021)
Cyst germination
Oospore production
A. fumigatus GP56 Major antigen/allergen with a molecular weight of 56 kD Studied only for virulence Nigam et al. (2003)
C. albicans CaHWP1 Hyphal wall protein 1 Positive regulators of hyphal development Sharkey et al. (1999); Staab et al. (2013)
CaALS1 Agglutinin-like sequence Studied only for virulence Hoyer et al. (2008); Kamai et al. (2002)
cell-surface glycoproteins
CaALS3 Agglutinin-like sequence Studied only for virulence Hoyer et al. (2008); Zhao et al. (2004)
cell-surface glycoproteins
CaALS2 Agglutinin-like sequence Studied only for virulence Hoyer et al. (2008); Zhao et al. (2005)
cell-surface glycoproteins
CaMP65 Putative β-glucanase mannoprotein Hyphal morphogenesis Sandini et al. (2007)
Fungal cell adherence
C. neoformans PLB1 Glycoproteins phospholipase Intracellular infection Evans et al. (2015); Thak et al. (2020)
Cell body morphology
MP65 Immunodominant 65-kDa-MP antigen Studied only for virulence Pietrella et al. (2002)
CIG1 Extracellular mannoprotein Iron acquisition Cadieux et al. (2013)
Growth on heme
MP84 Mannoprotein Not studied Teixeira et al. (2014)
CMP1 Cryptococcus mannoprotein 1 Capsule formation Han et al. (2020)
Localized in the vacuole
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        Integrative perspectives on glycosylation networks in fungi and oomycetes
        J. Microbiol. 2025;63(12):e2510003  Published online December 31, 2025
        DOI: https://doi.org/10.71150/jm.2510003
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      Figure
      Integrative perspectives on glycosylation networks in fungi and oomycetes
      Image Image
      Fig. 1. Summary of the pathogenic contributions of glycogenes associated with N- and O-glycosylation across representative fungal species. (A) Pathogenic contribution map of proteins involved in (A) N-glycosylation and (B) O-glycosylation pathways across fungal species. Species are color-coded; filled circles indicate genes with demonstrated pathogenic relevance, whereas open circles represent genes studied without pathogenic association. Abbreviations: Fg, Fusarium graminearum; Mo, Magnaporthe oryzae; Afu, Aspergillus fumigatus; Ca, Candida albicans; Cn, Cryptococcus neoformans. Filled circles indicate glycogenes that contribute to virulence, whereas unfilled circles represent glycogenes that have been characterized but do not affect virulence. Each positions indicate pathway step, not physical interaction.
      Fig. 2. Potential regulatory mechanisms of glycoproteins in fungal virulence. Functional analyses of glycogenes have been conducted in both human and plant pathogenic fungi. Conserved glycoproteins contribute to key processes such as protein maturation, cell wall integrity, morphogenesis, and virulence, yet their functional impacts vary in a species-specific manner across fungal lineages. (A) In plant pathogenic fungi, glycosylation regulates penetration and invasive growth by stabilizing and positioning key glycoproteins required for invasive growth, cell wall remodeling, immune evasion, and oxidative stress tolerance. These proteins can be grouped into ER quality control (ERQC; Cnx1, Gtb1, and Ecm33), autophagy and vesicle trafficking (Atg27, Syn2, Vps74, and Prb1), and cell wall integrity factors (Cwm1, Chs7, Chs5, and Arb1), all contributing to fungal development, sporulation, and mycotoxin production. Additional cell wall–localized or secretory proteins, such as Gpi7, KatG2, Glx, and LysM effectors (LysM1, Slp1, Scp1), contribute to invasive growth by scavenging host-derived reactive oxygen species and protecting fungal cell walls from plant chitinases. (B) In the human pathogen C. neoformans, several glycoproteins function as antigens and virulence determinants at the cell surface. Cig1, Mp65, and Mp84 act as immunogenic proteins that mediate host–pathogen interactions, while Cmp1 regulates capsule formation and Plb1 contributes to intracellular infection and morphological adaptation. Capsule polysaccharides further support immune evasion and pathogen survival. (C) In C. albicans, glycoproteins orchestrate adhesion, morphogenesis, and hyphal development. MP65 promotes fungal adhesion and morphogenetic transitions; Hwp1 facilitates hyphal development and stable adhesion to host tissues; and the Als family proteins (Als1, Als2, and Als3) function as major surface adhesins driving biofilm formation and immune evasion. This figure was created using BioRender.

      Fig. 1.

      Fig. 2.

      Integrative perspectives on glycosylation networks in fungi and oomycetes
      Species Gene Type Description Regulated genes Direct binding Reference
      C. albicans CaACE2 Zinc-finger RAM (regulation of ACE2 TF and polarized morphogenesis) signaling network PMTs Not studied Cantero and Ernst (2011); van Wijlick et al. (2016)
      B. cinerea BcFTG1 Zinc-finger fungal TF containing the GAL4 domain ALG8 Not studied Yang et al. (2024)
      H. polymorpha HpHAC1 bZIP Major unfolded protein response (UPR) ALG5, MNN2, MNN4, KTR1, OST1,OST2, OST4, OCR5, SWP1, WBP1 Not studied Moon et al. (2015)
      A. niger hacA bZIP Major unfolded protein response (UPR) ALG6, ALG8, ALG2, ALG3, DPM1,DPM2, ALG5, ALG7, RFT1, ALG9, SEC59, ALG12, ALG10, OST1, WBP1,OST2, OST3, SST3 Not studied Carvalho et al. (2012)
      C. neoformans HXL1 bZIP HAC1, XBP1-like gene 1 UPR genes (WBP1, PMT1, PMT2, PMT4, WBP1, ALG7, OST1) Not studied Cheon et al. (2011); Glazier et al. (2015)
      Mediate the unfolded protein response (UPR) pathway
      M. robertsii MrpacC C2H2-type zinc-finger pH-responsive TF STT3, OST3, PMT4 STT3, OST3, PMT4 Huang et al. (2015)
      M. acridum MaSte12 C2H2-type zinc-finger Direct target and functions downstream of the mitogen-activated protein kinase (MAPK) Fus3/Kss1 OCH1 Not studied Wei et al. (2017)
      Species Gene Virulence (O = affected; X = no effect) Biological function(s) Glycan structure Reference
      B. cinerea bcpmt1 O Morphogenesis Not studied González et al. (2013, 2014)
      Fungal adherence
      Cell wall integrity
      bcpmt2 O Morphogenesis Not studied González et al. (2013, 2014)
      Fungal adherence
      Cell wall integrity
      bcpmt4 O Morphogenesis Not studied González et al. (2013, 2014); Plaza et al. (2025)
      Fungal adherence
      Cell wall integrity
      C. graminicola Cgalg3 O ER homeostasis Not studied Mei et al. (2023)
      Fungal growth
      F. graminearum RFT1 O Mycotoxin synthesis Not studied Moon et al. (2025)
      Vegetative growth
      Conidiation
      Sexual development
      CWH41 (MOGS) X Mycotoxin synthesis Not studied
      Conidiation
      Sexual development
      ROT2 (GLS2, GANAB) X 6 orthologs showed different phenotypes Not studied
      Mycotoxin synthesis
      Vegetative growth
      Conidiation
      Sexual development
      ALG5 O Vegetative growth Not studied
      Sexual development
      ALG6 O Vegetative growth Not studied
      Sexual development
      ALG8 O Mycotoxin synthesis Not studied
      Vegetative growth
      Sexual development
      PSA1 (MPG) O Mycotoxin synthesis Not studied
      Vegetative growth
      Conidiation
      Sexual development
      MNS1 (MAN1B) O 2 orthologs showed different phenotypes Not studied
      Mycotoxin synthesis
      Conidiation
      MNL2 (MAN1A) O 4 orthologs showed different phenotypes Not studied
      Mycotoxin synthesis
      Vegetative growth
      Conidiation
      PMT1, 5, 7 O Vegetative growth Not studied
      Conidiation
      Sexual development
      MNN14 X Mycotoxin synthesis Not studied
      Vegetative growth
      Conidiation
      ALG1 X Vegetative growth Not studied
      ALG11 O Mycotoxin synthesis Not studied
      Vegetative growth
      Conidiation
      Sexual development
      ALG12 O Vegetative growth Biosynthesis of the Dol-PP-GlcNAc2Man8 (HPLC)
      Sexual development
      ALG2 o Mycotoxin synthesis Not studied
      Vegetative growth
      Conidiation
      Sexual development
      ALG3 O Mycotoxin synthesis Biosynthesis of the Dol-PP-GlcNAc2Man6 (HPLC)
      Vegetative growth
      Conidiation
      Sexual development
      ALG9 X Mycotoxin synthesis Not studied
      Vegetative growth
      HOC1 O Mycotoxin synthesis Not studied
      OCH1 O Mycotoxin synthesis Not studied
      Vegetative growth
      Conidiation
      KTR5, KTR7 O Mycotoxin synthesis Not studied
      Vegetative growth
      Conidiation
      Sexual development
      KTR4 O Vegetative growth Not studied
      Conidiation
      KRE2, KTR1, 2, 3, 6 X Mycotoxin synthesis Not studied
      Vegetative growth
      Sexual development
      MNT4 X Mycotoxin synthesis Not studied
      Conidiation
      MNN1, MNT3 X Mycotoxin synthesis Not studied
      MNN10 X Conidiation Not studied
      Sexual development
      MNN11 X Mycotoxin synthesis Not studied
      Vegetative growth
      Conidiation
      Sexual development
      MNN2, 5 X Conidiation Not studied
      MNN4 O Mycotoxin synthesis Not studied
      Vegetative growth
      VAN1 X Mycotoxin synthesis Not studied
      Vegetative growth
      Sexual development
      ANP1 O Mycotoxin synthesis Not studied
      Vegetative growth
      Conidiation
      Sexual development
      MNN9 O Vegetative growth Not studied
      Sexual development
      STT3 O Not changed Not studied
      SWP1 X Conidiation Not studied
      WBP1 O Mycotoxin synthesis Not studied
      Vegetative growth
      Conidiation
      F. oxysporum OCH1 O Fungal growth Not studied Li et al. (2014)
      Cell wall integrity
      hyphal adhesion
      GNT2 O Conidium size and morphology Biosynthesis of the Galactofuranose chains Lopez-Fernandez et al. (2015)
      Oxidative stress resistance (size-exclusion chromatography with Diffusion Ordered Spectroscopy (DOSY))
      cell wall organization, biogenesis and remodelling hyphal fusion
      Secretion of trafficking vesicles
      F. oxysporum f. sp. cucumerinum PMT1 O Fungal growth Not studied Li et al. (2014)
      Cell wall integrity
      Conidiation
      PMT2 O Fungal growth Not studied Li et al. (2014)
      Cell wall integrity
      Conidiation
      PMT4 O Fungal growth Not studied Xu et al. (2020b)
      Cell wall integrity
      Conidiation
      M. graminicola MgAlg2 O Cell wall integrity Biosynthesis of the (GlcNAc)2M3 Motteram et al. (2011)
      Yeast-like to filamentous growth switch (MALDI-ToF-MS)
      M. oryzae GLS1 O Mycelial growth Not studied Chen et al. (2020)
      Conidiation
      Invasive hyphal growth
      GLS2 O Mycelial growth Not studied Chen et al. (2020)
      Conidiation
      Invasive hyphal growth
      GTB1 O Mycelial growth Not studied Chen et al. (2020)
      Conidiation
      Invasive hyphal growth
      ALG3 O Mycelial growth Not studied Chen et al. (2014)
      Conidiation
      Invasive growth
      Oxidative stress resistance
      PMT2 O Fungal adhesion Not studied Guo et al. (2016)
      Conidial germination
      cell wall integrity
      Invasive hyphae growth
      PMT4 O Hyphal growth Not studied Pan et al. (2019)
      Conidiation
      Penetration and biotrophic invasion
      ALG9 O Conidial production Not studied Zhang et al. (2025)
      Appressorium formation
      Responses to stressors
      P. capsici PcSTT3B O Vegetative growth Not studied Cui et al. (2023)
      Sporangial release rate
      Zoospore production
      P. digitatum pdpmt2 O Cell wall integrity Not studied Harries et al. (2015)
      Conidiogenesis
      Sensitivity to fungicide
      U. maydis gls1 O Initial stages of biotrophic growth Not studied Fernández-Álvarez et al. (2013)
      gas1 O Intracellular expending Not studied Schirawski et al. (2005)
      gas2 O Intracellular expending Not studied Fernández-Álvarez et al. (2013)
      pmt4 O Appressorium formation Not studied Fernández-Álvarez et al. (2009)
      Plant cuticle penetration
      V. dahliae VdOCH1 O Hyphal growth Not studied Zhang et al. (2019b)
      Conidia production
      Microsclerotia formation
      Cell wall integrity
      VdSTT3 O Fungal development Not studied Su et al. (2018)
      Hyphal growth
      Glycoprotein secretion
      A. fumigatus CWH41 X Cell wall integrity (CWI) Not studied Zhang et al. (2008)
      Polarity
      Septation
      Conidiation
      AMS1 Not studied Conidial formation Not studied Li et al. (2009)
      Polarity
      Septation
      OCH1 O Sporulation Biosynthesis of the Outer glycan chain Kotz et al. (2010)
      (Capillary electrophoresis DNA Sequencer)
      PMT1 X No discernible phenotype Not studied Mouyna et al. (2010)
      ΔPMT1/PMT4 was lethal
      PMT2 O Essential Not studied Mouyna et al. (2010)
      PMT4 X Mycelial growth Not studied Mouyna et al. (2010)
      Conidiation
      Sensitivity to echinocandins
      ΔPMT1/PMT4 was lethal
      ANPA X Mycelial growth Core-mannan structure of fungal-type galactomannan (FTGM) Kadooka et al. (2022)
      Conidia formation (1H-NMR spectroscopy)
      MNN9 X No discernible phenotype Not involved in the synthesis of the FTGM α-core-mannan Du et al. (2019); Kadooka et al. (2022)
      Apical growth and polarity (1H-NMR spectroscopy)
      No significant role in the biosynthesis of N-glycans
      (MALDI-TOF MS)
      VAN1 X No discernible phenotype Not involved in the synthesis of the FTGM α-core-mannan Kadooka et al. (2022)
      (1H-NMR spectroscopy)
      KTR4 O Polarized growth Polymerization galactomannan Henry et al. (2019)
      Conidiation (HPLC analysis)
      Conidial viability
      KTR7 O Polarized growth Polymerization galactomannan Henry et al. (2019)
      Conidiation (HPLC analysis)
      Conidial viability
      CMSA Not studied Mycelial growth Biosynthesis of the FTGM core-mannan structure Onoue et al. (2018)
      sensitive to antifungal agents (Proton nuclear magnetic resonance (1H-NMR) spectroscopy)
      CMSB Not studied Mycelial growth Biosynthesis of the FTGM core-mannan structure Onoue et al. (2018)
      sensitive to antifungal agents (Proton nuclear magnetic resonance (1H-NMR) spectroscopy)
      MNT1 O Cell wall stability Not studied Wagener et al. (2008)
      MNN2 Not studied No discernible phenotype ΔMNN2/MNN5 Kadooka et al. (2024)
      ΔMNN2/MNN5 exhibited a growth defect and abnormal conidial formations Biosynthesis of the outer chain like α-(1→6)-linked mannan
      (1H-NMR analysis)
      MNN5 Not studied No discernible phenotype ΔMNN2/MNN5 Kadooka et al. (2024)
      ΔMNN2/MNN5 exhibited a growth defect and abnormal conidial formations Biosynthesis of the outer chain like α-(1→6)-linked mannan
      (1H-NMR analysis)
      STT3 Not studied Fungal growth Not studied Li et al. (2011)
      Cell wall integrity (CWI)
      GALF O Morphology and growth Biosynthesis of the GalfMan6GlcNAc2 Schmalhorst et al. (2008)
      Susceptible to drugs (Four-capillary electrophoresis DNA sequencer)
      A. nidulans ALG7 Not studied Failed to delete Not studied Gerhardt et al. (2025)
      ALG6 (ALGF) Not studied Not studied Not studied Anyaogu et al. (2021); Gerhardt et al. (2025)
      AMS1 X No visible effect on growth or morphology Not studied Anyaogu et al. (2021); Eades et al. (1998)
      Recycling macromolecular components
      AN5748 Not studied Fungal growth Not studied Gerhardt et al. (2025)
      Sporulation
      Resistance to sorbitol and tunicamycin
      PMTA Not studied Fungal growth Not studied Oka et al. (2004)
      Cell wall formation
      ΔPMTA/PMTC lethal
      PMTB Not studied Conidiation Not studied Goto et al. (2009)
      Polarity maintenance
      ΔPMTB/PMTC lethal
      PMTC Not studied Fungal growth Not studied Goto et al. (2009)
      Cell wall integrity
      Osmotic stabilization
      ΔPMTA/PMTC, ΔPMTB/PMTC lethal
      ALG1 Not studied Failed to delete Not studied Gerhardt et al. (2025)
      ALG2 Not studied Fungal growth Not studied Gerhardt et al. (2025)
      Sporulation
      ALG11 Not studied Failed to delete Not studied Gerhardt et al. (2025)
      ALG3 (ALGC) Not studied Sensitive to tunicamycin and calcium stress Biosynthesis of the Man5-7GlcNAc2 Anyaogu et al. (2021); Gerhardt et al. (2025)
      (UHPLC-FLR-MS)
      ALG9 (ALGI) Not studied Sensitive to tunicamycin and calcium stress Not studied Anyaogu et al. (2021); Gerhardt et al. (2025)
      ALG12 (ALGL) Not studied Failed to delete Not studied Anyaogu et al. (2021); Gerhardt et al. (2025)
      ALG13 Not studied Failed to delete Not studied Gerhardt et al. (2025)
      C. albicans CaCWH41 O Growth rates Not studied Mora-Montes et al. (2007)
      Cell wall composition
      CaROT2 o Growth rates Not studied Mora-Montes et al. (2007)
      Cell wall composition
      CaMNS1 O Growth rates Not studied Mora-Montes et al. (2007)
      Cell wall composition
      CaPMT2 O Essential Not studied Prill et al. (2005)
      CaPMT4 O Fungal growth Not studied Prill et al. (2005)
      Morphogenesis
      Antifungal resistance.
      CaPMT5 X Fungal growth Not studied Prill et al. (2005)
      Morphogenesis
      Antifungal resistance.
      CaPMT1 O (heterozygous)/ Hyphal morphogenesis Not studied Eades et al. (1998); Timpel et al. (2000)
      X (homozygous) Supersensitivity to the antifungal agents
      CaPMT6 O Morphogenesis Not studied Timpel et al. (2000)
      Antifungal sensitivities
      CaOCH1 O Temperature-sensitive growth Biosynthesis of the N-glycan outer chain Bates et al. (2006)
      Cellular aggregation (ES-MS analysis, Gas chromatography-mass spectrometry)
      Host-fungal interaction
      CaMNT1 O Adhesion Biosynthesis of the second mannose to O-glycan Buurman et al. (1998)
      (Biogel-P4 chromatography)
      CaMNN5 O Hyphal growth Biosynthesis of the N-linked mannan branches Bai et al. (2006)
      Iron homeostasis (Thin-layer chromatography and TLC plate to autoradiography)
      Cell wall integrity
      Morphogenesis
      CaMNT4 Not studied No discernible phenotype Not studied Mora-Montes et al. (2010)
      ΔMNT3/MNT4/MNT5 exhibited severe growth defect
      ΔMNT4/MNT5, ΔMNT3/MNT4/MNT5 altered cell wall composition
      CaMNT5 O (ΔMNT3/MNT5) No discernible phenotype Not studied Mora-Montes et al. (2010)
      ΔMNT3/MNT4/MNT5 exhibited severe growth defect
      ΔMNT4/MNT5, ΔMNT3/MNT4/MNT5 altered cell wall composition
      CaMNT3 O (ΔMNT3/MNT5) No discernible phenotype Not studied Mora-Montes et al. (2010)
      ΔMNT3/MNT5, ΔMNT3/MNT4/MNT5 exhibited severe growth defect
      ΔMNT3/MNT5, ΔMNT4/MNT5, ΔMNT3/MNT4/MNT5 altered cell wall composition
      C. neoformans MNS1 (ΔMNS1/MNS101) Sensitive to ER stress (tunicamycin, DTT) Remove mannose from GlcNAc2Man9 to GlcNAc2Man8 Mota et al. (2025)
      Sensitive to ER stress cell wall integrity stress (HPLC and MALDI-TOF)
      ΔMNS1/MNS101 growth defect
      MNS101 (ΔMNS1/MNS101) Sensitive to ER stress (tunicamycin, DTT) Remove mannose from (> M10) glycan or M8 further trimming Mota et al. (2025)
      Sensitive to ER stress cell wall integrity stress (HPLC and MALDI-TOF)
      ΔMNS1/MNS101 growth defect
      MNL1 X No discernible phenotype Involved in targeting misfolded proteins for ERAD Mota et al. (2025)
      rather than in normal N-glycan processing
      (HPLC, MALDI-TOF)
      MNL2 X No discernible phenotype Involved in targeting misfolded proteins for ERAD Mota et al. (2025)
      rather than in normal N-glycan processing
      (HPLC, MALDI-TOF)
      PMT4 O Fungal growth Not studied Olson et al. (2007); Willger et al. (2009)
      Cell wall integrity
      Cell morphology
      PMT2 O Essential Not studied Willger et al. (2009)
      PMT1 O Cell morphology and integrity Not studied Willger et al. (2009)
      ΔPMT1/PMT4 is synthetically lethal,
      ALG3 O Macrophage cell death Biosynthesis of the Dol-PP-GlcNAc2Man6 Thak et al. (2020)
      (HPLC, MALDI-TOF)
      OCH1 Slightly attenuated, No discernible phenotype Addition of a single α1,6-linked mannose residue to the Man8GlcNAc2 core Park et al. (2012)
      not critical (HPLC, MALDI-TOF)
      ALG9 O Sensitive to SDS, fluconazole Biosynthesis of the Dol-PP-GlcNAc2Man7 Thak et al. (2020)
      (HPLC, MALDI-TOF)
      MNN2 Not studied No discernible phenotype Biosynthesis of the elongated outer chain N-glycan Park et al. (2012)
      (HPLC, MALDI-TOF)
      KTR3 Not studied Cell wall stability Not involved in the processing of N-glycans (M6–10, X1M6–10) Park et al. (2012)
      (HPLC, MALDI-TOF)
      Species Gene Description Phenotypic defect(s) Reference
      B. cinerea bcsun1 Member of the β-glucosidase SUN family Fungal morphogenesis Pérez-Hernández et al. (2017)
      (secreted glycoprotein) Cell Wall Integrity
      Reproductive structures formation
      ROS sensitivity
      bcpg1 Endopolygalacturonase Studied only for virulence González et al. (2014); ten Have et al. (1998)
      bclysm1 Lysin motif domains Protects hyphae against degradation by external chitinases Crumière et al. (2025)
      Prevent them from inducing ROS
      Infection initiation
      Adhesion to host
      C. graminicola Cgcnx1 ER chaperone (Site directed mutagenesis) Mei et al. (2023)
      Vegetative growth
      Effector stability and secretion
      F. graminearum CFEMN1 Contain a CFEM domain Interact with extracellular binding Zuo et al. (2022)
      proteins from host
      TUB2 β2-tubulin Vegetative growth Liu et al. (2013);
      Sensitive to carbendazim Zhao et al. (2014)
      ADE5 Phosphoribosylamine-glycine ligase Adenine auxotrophy Kim et al. (2007)
      Vegetative growth
      Sexual reproduction
      CHS7 Chitin synthase Vegetative growth Kim et al. (2009)
      Septa formation
      Perithecia formation
      CHS5 Chitin synthase Vegetative growth Kim et al. (2009)
      Septa formation
      Perithecia formation
      VPS74 Vacuolar protein sorting Vegetative growth Kim et al. (2015b)
      Conidiation
      Sexual development
      Mycotoxin production
      SYN2 Syntaxin-like SNARE genes Self and female fertility Hong et al. (2010)
      Localized in plasma membranes and septa
      ERG9 Ergosterol biosynthesis Essential Yun et al. (2014)
      PRB1 Subtilisin-like protease belonging to proteinase K-like subfamily Mycotoxin production Xu et al. (2020a)
      Responses to environmental stimuli
      Lipid metabolism
      Autophagy regulation
      ATG27 Autophagy-related genes Vegetative growth Lv et al. (2017)
      Sporulation
      Mycotoxin production
      Autophagy regulation
      ILV6 Acetohydroxyacid synthase BCAA-auxotroph Liu et al. (2015)
      Vegetative growth
      Pigmentation
      Mycotoxin production
      ILV2 Acetohydroxyacid synthase BCAA-auxotroph Liu et al. (2015)
      Vegetative growth
      Conidial formation
      Pigmentation
      Mycotoxin production
      GPB1 G protein Mycotoxin production Yu et al. (2008)
      Chitin accumulation
      RAB11 Ras-related proteins in brain Polarized growth and/or exocytosis Zheng et al. (2015)
      KAR Hsp70s Essential Liu et al. (2017)
      SSC Hsp70s Essential Liu et al. (2017)
      SSB Hsp70s Proper folding of β2-tubulin Liu et al. (2017)
      Vegetative growth
      Low temperature response
      Vacuole fusion and endocytosis
      Mycotoxin production
      SSA Hsp70s Essential Liu et al. (2017)
      CAP1 Cyclase-associated protein Vegetative growth Yin et al. (2018b)
      Conidiogenesis
      Mycotoxin production
      KATG2 /FCA7 peroxidase (Site directed mutagenesis) Guo et al. (2019);
      Scavenges ROS Lee et al. (2018)
      Located on the cell wall of invading hyphal cells
      ECM33 GPI-anchored protein Cell wall morphology Fernando et al. (2019)
      PP2A Phosphatase Essential Yu et al. (2014)
      STE7 kinase Vegetative growth Gu et al. (2015)
      Conidiation
      Mycotoxin production
      Penetration
      PHO8 Phosphatase Essential Yun et al. (2015)
      CNA1 Phosphatase Essential Yun et al. (2015)
      Fg07304 Phosphatase Essential Yun et al. (2015)
      GIC1 Phosphatase Conidiation Yun et al. (2015)
      PSR2 Phosphatase Vegetative growth Yun et al. (2015)
      Pigmentation
      Conidia morphology
      ARC15 Phosphatase Only defects in virulence Yun et al. (2015)
      GLX glyoxal oxidase Mycotoxin production Song et al. (2016)
      localized to cell membrane
      ARB1 ATP-binding protein Infective growth Yin et al. (2018a)
      Oxidative stresses
      Cell wall integrity
      Mycotoxin production
      Sexual and asexual development
      SKP1 E3 ubiquitin ligases Essential Chen et al. (2023)
      LIP50, TAG Lipases Only defects in virulence Kim et al. (2023); Zhang et al. (2016)
      PCT1 Cytidylyltransferases Essential Wang et al. (2019)
      FGSG_10825 Mating-type Sexual development Kim et al. (2015a)
      OB031, HEX1 TF Only defects in virulence Son et al. (2011)
      CMK1 Kinase Ascospore formation Wang et al. (2011)
      ALDH (FGSG_02273) Aldehyde Dehydrogenase Essential Tang et al. (2023)
      ALDH (FGSG_04194) Aldehyde Dehydrogenase Regulating acetaldehyde dehydrogenase activity Tang et al. (2023)
      Accumulation of lipid droplets
      Conidial germination
      CWM1 Cell wall mannoprotein Vegetative growth Zhang et al. (2019a)
      Cell wall integrity
      L. theobromae LtScp1 Effectors, a LysM-containing protein (Site directed mutagenesis) Peng et al. (2022)
      Interacted with chitinase VvChi4
      M. oryzae CNX1 Calnexin Mycelial growth Chen et al. (2020)
      Conidiation
      Invasive hyphal growth
      GTB1 Glucosidase II β-subunit Gas2 Mycelial growth Chen et al. (2020)
      Conidiation
      Invasive hyphal growth
      SLP1 Secreted LysM domain protein (Site directed mutagenesis) Chen et al. (2014); Jones et al. (2021)
      Sequesters chitin oligosaccharides
      Apoplastic effectors
      BAS4 Biotrophy-associated secreted protein 4 Apoplastic effectors Chen et al. (2014); Jones et al. (2021)
      GPI7 Glycosylphosphatidylinositol Penetration and invasive growth Liu et al. (2020)
      Cell wall biogenesis
      P. capsici Pcipg2 Polygalacturonase Studied only for virulence Sun et al. (2009)
      P. sojae GPI16 GPI transamidase component protein (Site directed mutagenesis) Zhang et al. (2021)
      Cyst germination
      Oospore production
      A. fumigatus GP56 Major antigen/allergen with a molecular weight of 56 kD Studied only for virulence Nigam et al. (2003)
      C. albicans CaHWP1 Hyphal wall protein 1 Positive regulators of hyphal development Sharkey et al. (1999); Staab et al. (2013)
      CaALS1 Agglutinin-like sequence Studied only for virulence Hoyer et al. (2008); Kamai et al. (2002)
      cell-surface glycoproteins
      CaALS3 Agglutinin-like sequence Studied only for virulence Hoyer et al. (2008); Zhao et al. (2004)
      cell-surface glycoproteins
      CaALS2 Agglutinin-like sequence Studied only for virulence Hoyer et al. (2008); Zhao et al. (2005)
      cell-surface glycoproteins
      CaMP65 Putative β-glucanase mannoprotein Hyphal morphogenesis Sandini et al. (2007)
      Fungal cell adherence
      C. neoformans PLB1 Glycoproteins phospholipase Intracellular infection Evans et al. (2015); Thak et al. (2020)
      Cell body morphology
      MP65 Immunodominant 65-kDa-MP antigen Studied only for virulence Pietrella et al. (2002)
      CIG1 Extracellular mannoprotein Iron acquisition Cadieux et al. (2013)
      Growth on heme
      MP84 Mannoprotein Not studied Teixeira et al. (2014)
      CMP1 Cryptococcus mannoprotein 1 Capsule formation Han et al. (2020)
      Localized in the vacuole
      Table 1. TFs regulating glycogenes

      Table 2. N&O-Glycogenes and glycan structures discussed in this review

      Man9GlcNAc2 : Glycan compositions with 9 mannose and two N-acetylglucosamine residues.

      X1M6–10: Extended high-mannose glycans containing one Xylose and 6–10 mannose residues.

      FTGM: Fungal-type galactomannan.

      Table 3. Glycoproteins in fungal virulence and development

      Table 1.

      Table 2.

      Table 3.

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