Exploring fern pathosystems and immune receptors to bridge gaps in plant immunity
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Abstract
Land plants include angiosperms, gymnosperms, bryophytes, lycophytes, and ferns, each of which may deploy distinct strategies to resist pathogens. Here, we investigate fern-pathogen interactions by characterizing novel pathosystems and analyzing the diversity of fern immune receptors. A collection of fern species was inoculated with a diverse set of filamentous microbes, and disease symptoms were assessed. We further leveraged published genome mining tools to analyse the diversity of receptor-like kinases, receptor-like proteins (RLKs/RLPs) and nucleotide-binding and leucine-rich repeats (NLRs), along with key immune signalling components, in ferns. Our results reveal that ferns exhibit a range of responses to pathogens, including putative non-host resistance and more specific resistance mechanisms. Among ten ferns tested, Pteris vittata displays the broadest spectrum of pathogen compatibility. Genome mining indicates that ferns encode a diverse repertoire of putative immune receptors, antimicrobial peptides, and mediators of systemic acquired resistance. Ferns possess numerous RLKs/RLPs, resembling those required for cell-surface immunity in angiosperms. They also encode diverse NLRs, including sub-families lost in flowering plants. These findings provide insights into disease resistance evolution and open promising perspectives for crop protection strategies.
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Moussart, A., Onfroy, C., Lesné, A., Esquibet, M., Grenier, E., & Tivoli, B. (2006) · European journal of plant pathology · doi:10.1007/s10658-006-9071-y · Cited by 54
The evolution of plant NLR immune receptors and downstream signal components.
Liu, Y., Zhang, Y.-M., Tang, Y., Chen, J.-Q., & Shao, Z.-Q. (2023) · Current opinion in plant biology · doi:10.1016/j.pbi.2023.102363 · Cited by 49 · cited 1×
“Bryophytes additionally contain lineage-specific kinase NLRs (Kin-NLRs) and α/β-hydrolase NLRs (Hyd-NLRs) [24–26].”
Conservation of the PBL-RBOH immune module in land plants.
Chu, J., Monte, I., DeFalco, T. A., Köster, P., Derbyshire, P., Menke, F., & Zipfel, C. (2023) · Current Biology · doi:10.1016/j.cub.2023.01.050 · Cited by 48 · cited 1×
“This is corroborated by functional studies showing conservation of central PTI regulators such as LysM receptors [112, 113], SERK receptors [114], CPK28 [115], or RBOH [116].”
Redkar, A., Ibanez, S. G., Sabale, M., Zechmann, B., Solano, R., & Pietro, A. D. (2021) · bioRxiv · doi:10.1101/2021.03.20.436100 · Cited by 48
Encyclopedia of Plant Viruses and Viroids
Sastry, K., Mandal, B., Hammond, J., Scott, S., & Briddon, R. (2019) · doi:10.1007/978-81-322-3912-3 · Cited by 46 · cited 1×
“Yet, several pathogens have been reported on ferns, primarily filamentous fungi [16, 74– 76], but also bacteria [77–79] and viruses [80].”
Quantitative interactions: the disease outcome of Botrytis cinerea across the plant kingdom
Caseys, C., Shi, G.-J., Soltis, N., Gwinner, R., Corwin, J. A., Atwell, S., & Kliebenstein, D. (2021) · G3 · doi:10.1093/g3journal/jkab175 · Cited by 43
The ecology and physiology of fern gametophytes: A methodological synthesis
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From Gene-for-Gene to Resistosomes: Flor's Enduring Legacy.
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“Another model is the gene-for-gene concept, in which a single molecular factor from the pathogen is recognized by a single molecular factor in the plant, triggering immunity [20].”
Kloepper, J., McInroy, J. A., Liu, K. X., & Hu, C.-H. (2013) · PLoS ONE · doi:10.1371/journal.pone.0058531 · Cited by 37 · cited 1×
“Yet, several pathogens have been reported on ferns, primarily filamentous fungi [16, 74– 76], but also bacteria [77–79] and viruses [80].”
Tausz, M., Hietz, P., & Briones, O. (2001) · doi:10.1071/pp01068 · Cited by 36 · cited 1×
“Fern gameto-phytes also show adaptation to light stress [94], likely through the accumulation of reactive oxygen species-scavenging molecules, such as β-carotene or tocophe-rols [95, 96].”
Lehti-Shiu, M., Zou, C., & Shiu, S. (2012) · Signaling and communication in plants · doi:10.1007/978-3-642-23044-8_1 · Cited by 35 · cited 1× · see citing works
“, wall-associated kinase (WAK), thaumatin, cysteine-rich repeat (CRR), regulator of chromosome condensation 1 (RCC1), and lysin motif (LysM) (Additional file1: Table S2) [29, 60].”
Chia, K.-S., Kourelis, J., Teulet, A., Vickers, M. J., Sakai, T., Walker, J. F., Schornack, S., Kamoun, S., & Carella, P. (2024) · The Plant Cell · doi:10.1093/plcell/koae113 · Cited by 34 · see citing works
Johanndrees, O., Baggs, E., Uhlmann, C., Locci, F., Läßle, H. L., Melkonian, K., Käufer, K., Dongus, J. A., Nakagami, H., Krasileva, K. V., et al. (2022) · Plant Physiology · doi:10.1093/plphys/kiac480 · Cited by 32 · cited 1× · see citing works
“TIR-NB-TPRs form a conserved class of NLRs, potentially regulating immunity, containing a TIR domain but signalling independently of TIR-NLRs [61].”
Yotsui, I., Matsui, H., Miyauchi, S., Iwakawa, H., Melkonian, K., Schlüter, T., Michavila, S., Kanazawa, T., Nomura, Y., Stolze, S., et al. (2023) · Current Biology · doi:10.1016/j.cub.2023.07.068 · Cited by 31
Anti-microbial Activity of Phenolic Acids in Pteridium aquilinum
Francisco, M. S., & Cooper-Driver, G. (1984) · doi:10.2307/1546543 · Cited by 31
Алексеевна, К. О., Kliukova, M. S., Afonin, A. М., Sulima, A. S., Zhukov, V. A., & Tikhonovich, I. A. (2020) · Biological Communications · doi:10.21638/spbu03.2020.205 · Cited by 30 · see citing works
TIR-domain enzymatic activities at the heart of plant immunity.
Locci, F., Wang, J.-L., & Parker, J. (2023) · Current opinion in plant biology · doi:10.1016/j.pbi.2023.102373 · Cited by 28 · cited 2× · see citing works
“They have been shown to function in the same immune signalling pathway, along with the protein EDS1 [107, 120, 121].”
Freezing resistance of gametophytes of the temperate fern, Polystichum retroso-paleaceum
Sato, T., & Sakai, A. (1980) · doi:10.1139/b80-141 · Cited by 28 · cited 1×
“Additionally, some fern gametophytes are more resistant to freezing than their sporophytes, enabling their survival under snow during winter [97].”
Ghosh, Mukhopadhyay, R., & Gupta, K. (2005) · doi:10.6165/tai.2005.50(4).272 · Cited by 27
Li, C., Wickell, D., Kuo, L., Chen, X., Nie, B., Liao, X.-Z., Peng, D., Ji, J.-J., Jenkins, J., Williams, M., et al. (2023) · bioRxiv · doi:10.1073/pnas.2312607121 · Cited by 26 · see citing works
Contrasting and conserved roles of NPR pathways in diverged land plant lineages
Jeon, H.-W., Iwakawa, H., Naramoto, S., Herrfurth, C., Gutsche, N., Schlüter, T., Kyozuka, J., Miyauchi, S., Feussner, I., Zachgo, S., et al. (2023) · bioRxiv · doi:10.1101/2022.07.19.500630 · Cited by 26 · cited 1×
“In Marchantia polymorpha , NPR1 is conserved, can complement Arabidopsis npr1 mutants, and can bind SA, suggesting that this function is ancestral to land plants [108].”
Wang, Y.-Q., Teng, Z., Li, H., Wang, W., Xu, F., Sun, K., Chu, J.-F., Qian, Y.-W., Loake, G., Chu, C., et al. (2022) · Plant Communications · doi:10.1016/j.xplc.2022.100459 · Cited by 26 · cited 1×
“The rice CC-NB-ARM “RLS1” could be a regulator of immunity [64].”
Genomic Insights into Genetic Diploidization in the Homosporous Fern Adiantum nelumboides
Zhong, Y., Liu, Y.-B., Wu, W., Chen, J.-F., Sun, C.-Y., Liu, H., Shu, J.-P., Ebihara, A., Yan, Y.-H., Zhou, R., et al. (2022) · Genome Biology and Evolution · doi:10.1093/gbe/evac127 · Cited by 26 · cited 2× · see citing works
“…Selaginella moellendorffii [52]), ferns ( Alsophila spinulosa [9], Ceratopteris richardii [6], Adiantum capillus-veneris [7], Adiantum nelumboides [8], Azolla filiculoides [4], Salvinia cucullata [4], Mar-silea vestita [5]); monocots ( Dendrobium catenatum [53], Oryza sativa [54], Setaria italica…”
Goh, F.-J., Huang, C.-Y., Derevnina, L., & Wu, C.-H. (2024) · bioRxiv · doi:10.1101/2023.10.25.563953 · Cited by 25
Johnson, D., Carris, L. M., & Rogers, J. (1997) · Mycological Research · doi:10.1017/s0953756296003115 · Cited by 24 · cited 1× · see citing works
“C. nymphaeae produced appressorium-like structures, suggesting a penetration strategy similar to that on waterlil-ies and strawberries [42, 43].”
NLR receptor networks in plants.
Adachi, H., & Kamoun, S. (2022) · Essays in Biochemistry · doi:10.1042/ebc20210075 · Cited by 23 · cited 1×
“Their success in asterids could be explained by the robustness of the “NRC” networks, formed between many CC-NLRs “EDVID” [105, 117–119].”
Rahmatpour, N., Kuo, L., Kang, J., Herman, E., Lei, L., Li, M., Srinivasan, S., Zipper, R. S., Wolniak, S. M., Delwiche, C., et al. (2022) · New Phytologist · doi:10.1111/nph.18652 · Cited by 20
Ali, Z., Tan, Q., Lim, P.-K., Chen, H.-H., Pfeifer, L., Julca, I., Lee, J.-M., Classen, B., de Vries, S., de Vries, J., et al. (2025) · Nature Plants · doi:10.1038/s41477-025-01978-y · Cited by 19 · see citing works
Immunobiodiversity: conserved and specific immunity across land plants and beyond.
Castel, B., Mahboubi, K. E., Jacquet, C., & Delaux, P. (2023) · Molecular Plant · doi:10.1016/j.molp.2023.12.005 · Cited by 18 · cited 2× · see citing works
“Bryophytes additionally contain lineage-specific kinase NLRs (Kin-NLRs) and α/β-hydrolase NLRs (Hyd-NLRs) [24–26].”
Self-citationFurumizu, C., & Sawa, S. (2021) · Plant Molecular Biology · doi:10.1007/s11103-020-01100-0 · Cited by 18 · cited 1×
“Fern genomes encode RLKs, including families that are well described in angiosperms such as LRR-RLKs and LysM-RLKs [29, 30].”
Equisetum species show uniform epicuticular wax structures but diverse composition patterns
Brune, T., & Haas, K. (2011) · AoB Plants · doi:10.1093/aobpla/plr009 · Cited by 18 · cited 1×
“For example, they have a distinct cuticle wax layer, known to contribute to resilience against abiotic stress [85], which could also play a role in disease resistance.”
A necrotizing toxin enables Pseudomonas syringae infection across evolutionarily divergent plants.
Grenz, K., Chia, K.-S., Turley, E., Tyszka, A. S., Atkinson, R. E., Reeves, J., Vickers, M. J., Rejžek, M., Walker, J. F., & Carella, P. (2024) · Cell Host and Microbe · doi:10.1016/j.chom.2024.11.014 · Cited by 17 · see citing works
A fungal plant pathogen discovered in the Devonian Rhynie Chert
Strullu-Derrien, C., Góral, T., Spencer, A. R. T., Kenrick, P., Aime, M., Gaya, E., & Hawksworth, D. L. (2023) · Nature Communications · doi:10.1038/s41467-023-43276-1 · Cited by 16 · cited 1×
“Tracheophytes have been exposed to pathogen infections for at least 407 million years [104] and likely since terres-trialization [26, 105, 106].”
CorkOakDB—The Cork Oak Genome Database Portal
Arias-Baldrich, C., Silva, M., Bergeretti, F., Chaves, I., Miguel, C. M., Saibo, N. J. M., Sobral, D., Faria, D., & Barros, P. (2020) · Database J. Biol. Databases Curation · doi:10.1093/database/baaa114 · Cited by 16 · cited 1×
“…[7], Adiantum nelumboides [8], Azolla filiculoides [4], Salvinia cucullata [4], Mar-silea vestita [5]); monocots ( Dendrobium catenatum [53], Oryza sativa [54], Setaria italica [55]); and dicots ( Camellia sinensis [56], Arabidopsis thaliana [57], Medicago truncatula [58], Quercus suber [59]).”
Molecular perspectives on age-related resistance of plants to (viral) pathogens.
DeMell, A., Alvarado, V., & Scholthof, H. (2023) · New Phytologist · doi:10.1111/nph.19131 · Cited by 15 · cited 1×
“In angiosperms, developmental stage-dependent resistance to pathogens has been also reported, with mature plants generally more resistant than seedlings [98, 99].”
Host and nonhost resistance in Medicago-Colletotrichum interactions.
Jaulneau, V., Cazaux, M., Hoi, J. W. S., Fournier, S., Esquerré-Tugayé, M., Jacquet, C., & Dumas, B. (2010) · Molecular Plant-Microbe Interactions · doi:10.1094/mpmi-23-9-1107 · Cited by 15
Self-citationDo fern gametophytes have the capacity for irradiance acclimation?
Fernández-Marín, B., Alfaro, S. J. A., Becerril, J., & García-Plazaola, J. (2012) · Biologia plantarum · doi:10.1007/s10535-012-0097-z · Cited by 13 · cited 1×
“Fern gameto-phytes also show adaptation to light stress [94], likely through the accumulation of reactive oxygen species-scavenging molecules, such as β-carotene or tocophe-rols [95, 96].”
A plant Lysin Motif Receptor-Like Kinase plays an ancestral function in mycorrhiza
Teyssier, E., Grat, S., Landry, D., Ouradou, M., Rich, M., Fort, S., Keller, J., Lefebvre, B., Delaux, P., & Mbengue, M. (2025) · Proceedings of the National Academy of Sciences of the United States of America · doi:10.1073/pnas.2426063122 · Cited by 11
Self-citationMartinez, A., Chiocchio, V., Em, L., Rodríguez, M. A., & Godeas, A. (2012) · Revista de Biología Tropical · doi:10.15517/rbt.v60i2.4019 · Cited by 11 · see citing works
A Sketch of the History of Fern Classification
Tryon, R. (1952) · doi:10.2307/2399092 · Cited by 10 · cited 1×
“Historically, ferns have been primarily studied through morphological traits [10, 11].”
Leaf spot and blight of Asplenium nidus caused by Pseudomonas gladioli
Chase, A., Miller, J. W., & Jones, J. B. (1984) · doi:10.1094/pd-69-344 · Cited by 10 · cited 1×
“Yet, several pathogens have been reported on ferns, primarily filamentous fungi [16, 74– 76], but also bacteria [77–79] and viruses [80].”
Cui, J.-T., Zhu, Y., Du, H., Liu, Z., Shen, S.-Q., Wang, T., Cui, W., Zhang, R., Jiang, S., Wu, Y.-M., et al. (2022) · GigaScience · doi:10.1093/gigascience/giad079 · Cited by 9 · cited 1×
“…Marchantia polymorpha [47], Physcomitrium patens [48], Takakia lepidozioides [49]); lycophytes ( Diphasiastrum complanatum [50], Isoetes sinensis [51], Selaginella moellendorffii [52]), ferns ( Alsophila spinulosa [9], Ceratopteris richardii [6], Adiantum capillus-veneris [7], Adiantum…”
Knowlton, A. A. (2012) · Current Biology · doi:10.1016/j.cub.2012.04.005 · Cited by 9 · cited 1×
“…pattern, ETI: effector‑triggered immunity, CNL: CC‑NLR, TNL: TIR‑NLR, SAR: systemic acquired resistance, AMP: antimicrobial peptide taken over the photosynthetic function of the leaf [84], they remain structurally different from true leaves, which could hinder pathogen infection in our experiment.”
Diversity of Colletotrichum species on strawberry (Fragaria × ananassa) in Germany
Rose, C., & Damm, U. (2024) · Phytopathologia Mediterranea · doi:10.36253/phyto-15094 · Cited by 8
Kim, H., Prokchorchik, M., & Sohn, K. (2022) · The Plant Journal · doi:10.1111/tpj.15653 · Cited by 8
Phylogenomic insights into the diversity and evolution of RPW8-NLRs and their partners in plants.
Qin, H., Cheng, J., Han, G.-Z., & Gong, Z. (2024) · The Plant Journal · doi:10.1111/tpj.17034 · Cited by 7 · cited 3× · see citing works
“TIR-NLRs and RPW8-NLRs are found in all lineages (Additional file1:Table S3) [67].”
InfluentialConserved role of the SERK-BIR module in development and immunity across land plants.
Yan, Y., Mellüh, J., Mecchia, M., Jeon, H.-W., Melkonian, K., Holzberger, C., Harzen, A., Stolze, S., Neuman, U., Franzen, R., et al. (2025) · Current Biology · doi:10.1016/j.cub.2025.03.072 · Cited by 6 · cited 1× · see citing works
“This is corroborated by functional studies showing conservation of central PTI regulators such as LysM receptors [112, 113], SERK receptors [114], CPK28 [115], or RBOH [116].”
Pathogenic Fungi in Ferns and Angiosperms: A Comparative Study
Antonovics, J. (2020) · American Fern Journal · doi:10.1640/0002-8444-110.3.79 · Cited by 6 · see citing works
Stevenson, J. (1945) · doi:10.2307/1545645 · Cited by 6 · cited 1×
“Yet, several pathogens have been reported on ferns, primarily filamentous fungi [16, 74– 76], but also bacteria [77–79] and viruses [80].”
Jones, K., Rayamajhi, M., Pratt, P., & Van, T. (2003) · Plant Disease · doi:10.1094/pdis.2003.87.1.101b · Cited by 6 · cited 2×
“Some of the reported filamentous pathogens belong to the same species as crop pathogens, such as Colletotrichum gloeosporioides [14] and Botrytis cinerea [16], both causing diseases on many crops including soybean, wheat, and tomato [81, 82].”
Mitoi, E., Aldea, F., Helepciuc, F., Ciocan, A., Frum, A., Popescu, D., Luțu, O., Șuțan, N. A., & Soare, L. (2024) · Horticulturae · doi:10.3390/horticulturae10080799 · Cited by 5 · see citing works
Mali, A., Patil, V., Pise, N., & Ade, A. (2016) · doi:10.1094/pdis-04-15-0427-pdn · Cited by 3 · cited 1×
“In the USA alone, at least 746 filamentous pathogens have been reported on ferns, including ascomycetes such as Colletotrichum [14] and Fusarium species [15], basidi-omycetes such as Puccinia and Hyalopsora species, and oomycetes such as Pythium and Achlya species [16].”
Divergent roles of CPK28 in immune homeostasis across land plants
Dou, R., Castel, B., Tanney, C. A. S., Bredow, M., Miguel, V. N., Mahboubi, K. E., Chu, J., Melkonian, K., Gallo, M. C. R., Lauressergues, D., et al. (2026) · bioRxiv · doi:10.1101/2024.10.01.616128 · Cited by 2 · cited 1×
“This is corroborated by functional studies showing conservation of central PTI regulators such as LysM receptors [112, 113], SERK receptors [114], CPK28 [115], or RBOH [116].”
Self-citationFern Family Clade Age and Fungal Pathogen Diversity
Antonovics, J. (2023) · American Fern Journal · doi:10.1640/0002-8444-113.3.137 · Cited by 2 · cited 3×
“Some of the reported filamentous pathogens belong to the same species as crop pathogens, such as Colletotrichum gloeosporioides [14] and Botrytis cinerea [16], both causing diseases on many crops including soybean, wheat, and tomato [81, 82].”
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Molecular mechanisms of plant NLR activation and signalling
Maruta, N., Sorbello, M., Garzón-Flores, L., & Kobe, B. (2026) · The Plant Journal · doi:10.1111/tpj.70702 · Cited by 11
Variations on a theme: Non-canonical mechanisms of effector-triggered immunity.
Ahn, H.-K., Jones, J. D. G., & Guo, G.-H. (2025) · Current opinion in plant biology · doi:10.1016/j.pbi.2025.102812 · Cited by 4
Longevity stabilizes core gene families and reshapes NLR architecture in trees
Fang, Y.-P. (2026) · Forestry Research Advances · doi:10.48130/fra-0026-0017
- Authors
- Castel, B., Baker, M., Keller, J., Martinez, Y., Bonhomme, M., Delaux, P.-M., & Jacquet, C.
- Affiliations
- Centre National de la Recherche Scientifique (FR)
- Fédération de Recherche Agrobiosciences, Interactions et Biodiversité (FR)
- Corresponding author
- Jacquet, Christophe
- ORCID
- Venue
- BMC Biology
- Published
- 2025-10-09
- Volume
- 23
- Issue
- 1
- Pages
- 301-301
- ISSN
- 1741-7007 (ISSN-L)
- PMC
- PMC12512272
- OpenAlex
- W4414985227
- Cited by
- 4
- References
- 128
- Citations by year
- 2026: 3 · 2025: 1
- FWCI
- 2.15
- Type
- journal article
- Language
- en
- Open access
- gold
- doi.org
- Locations
- BMC Biology (publishedVersion, cc-by)
- bioRxiv (Cold Spring Harbor Laboratory) (acceptedVersion, cc-by-nc)
- Europe PMC (PubMed Central) (publishedVersion, cc-by)
- PubMed (publishedVersion)
- HAL (submittedVersion, cc-by)
- DOAJ (DOAJ: Directory of Open Access Journals) (submittedVersion, cc-by-sa)
- Funders
- Agence Nationale de la Recherche (ANR-21-CE20-0010-01)
- HORIZON EUROPE European Research Council (101001675)
- Ministère de l'Education Nationale, de l'Enseignement Supérieur et de la Recherche
- Topics
- Life Sciences › Agricultural and Biological Sciences › Plant Science › Plant-Microbe Interactions and Immunity
- Life Sciences › Agricultural and Biological Sciences › Plant Science › Plant Disease Resistance and Genetics
- Life Sciences › Agricultural and Biological Sciences › Plant Science › Plant Pathogenic Bacteria Studies
- Keywords
- ferns, plant immunity, immune receptors, receptor-like kinases, receptor-like proteins, Pteris vittata, nonhost resistance, systemic acquired resistance, antimicrobial peptides, disease resistance evolution, genome mining
- MeSH terms
- Plant Diseases, Plant Proteins, Receptors, Immunologic, Ferns, Host-Pathogen Interactions, Plant Immunity, Disease Resistance
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