Abstract
Ectomycorrhizal fungi (EMF) form symbiotic relationships with approximately 60% of trees and woody plants among temperate and boreal forests. EMF also serve as the primary symbionts in subtropical and tropical forests, particularly in conifer forests, including those in Florida's climate. These fungi influence the global carbon, nitrogen, and phosphorus cycles, sequestering or supplying these elements as necessary for ecosystem processes and resilience. However, EMF are susceptible to environmental stressors and management practices, including fertilizers, fungicides, and mechanical disturbances. This publication aims to provide general knowledge to the general public on the biology and ecological functions of EMF in natural and managed forest ecosystems. By understanding the role and benefits of EMF in these environments, forest managers, tree nursery operators, and timber industry stakeholders can gain valuable insights for incorporating EMF into their management practices.
References
Alvarez, M., D. Huygens, C. Fernandez, et al. 2009. “Effect of Ectomycorrhizal Colonization and Drought on Reactive Oxygen Species Metabolism of Nothofagus dombeyi Roots.” Tree Physiology 29 (8): 1047–1057. https://doi.org/10.1093/treephys/tpp038
Andres, H., H.-L. Liao, and K. Zhang. 2025. “Biology, Ecology, and Benefits of Arbuscular Mycorrhizal Fungi in Agricultural Ecosystems: PP383, 3/2025.” EDIS 2025 (2). https://doi.org/10.32473/edis-pp383-2025
Anthony, M. A., T. W. Crowther, S. van der Linde, et al. 2022. “Forest tree growth is linked to mycorrhizal fungal composition and function across Europe.” The ISME Journal 16 (5): 1327–1336. https://doi.org/10.1038/s41396-021-01159-7
Ainsworth, E. A., and S. P. Long. 2005. “What have we learned from 15 years of free‐air CO2 enrichment (FACE)? A meta‐analytic review of the responses of photosynthesis, canopy properties and plant production to rising CO2.” New Phytologist 165 (2): 351–372. https://doi.org/10.1111/j.1469-8137.2004.01224.x
Aryal, P., S. J. Meiners, and B. S. Carlsward. 2021. “Ectomycorrhizae determine chestnut seedling growth and drought response.” Agroforestry Systems 95: 1251–1260. https://doi.org/10.1007/s10457-020-00488-4
Assad, R., Z. A. Reshi, I. Rashid, and Y. Shouche. 2020. “Role of Ectomycorrhizal Biotechnology in Pesticide Remediation.” Bioremediation and Biotechnology 3: 315–330. Springer Nature. https://doi.org/10.1007/978-3-030-46075-4_14
Bandou, E., F. Lebailly, F. Muller, et al. 2006. “The ectomycorrhizal fungus Scleroderma bermudense alleviates salt stress in seagrape (Coccoloba uvifera L.) seedlings.” Mycorrhiza 16: 559–565. https://doi.org/10.1007/s00572-006-0073-6
Baslam, M., I. Garmendia, and N. Goichoechea. 2011. “Arbuscular mycorrhizal fungi (AMF) improved growth and nutritional quality of greenhouse-grown lettuce.” Journal of Agricultural and Food Chemistry 59 (10): 5504–5515. https://doi.org/10.1021/jf200501c
Bauman, J. M., C. H. Keiffer, and S. Hiremath. 2012. “Facilitation of American Chestnut (Castanea dentata) Seedling Establishment by Pinus virginiana in Mine Restoration.” International Journal of Ecology 2012: 257326. https://research.fs.usda.gov/treesearch/41681
Begum, N., C. Qin, M. A. Ahanger, et al. 2019. “Role of Arbuscular Mycorrhizal Fungi in Plant Growth Regulation: Implications in Abiotic Stress Tolerance.” Frontiers in Plant Science 10: 1068. https://doi.org/10.3389/fpls.2019.01068
Berrios, L., G. D. Bogar, A. M. Venturini, et al. 2024. “Ectomycorrhizal fungi alter soil food webs and the functional potential of bacterial communities.” ASM Journals 9 (6): e00369-24. https://doi.org/10.1128/msystems.00369-24
Birhane, E., F. Sterck, M. Fetene, F. Bongers, and T. Kuyper. 2012. “Arbuscular mycorrhizal fungi enhance photosynthesis, water use efficiency, and growth of frankincense seedlings under pulsed water availability conditions.” Oecologia 169: 895–904. https://doi.org/10.1007/s00442-012-2258-3
Bradford, M. A., N. Fierer, and J. F. Reynolds. 2008. “Soil carbon stocks in experimental mesocosms are dependent on the rate of labile carbon, nitrogen and phosphorus inputs to soils.” Functional Ecology 22 (6): 964–974. https://doi.org/10.1111/j.1365-2435.2008.01404.x
Branco, S., A. Schauster, H.-L. Liao, and J. Ruytinx. 2022. “Mechanisms of Stress Tolerance and Their Effects on the Ecology and Evolution of Mycorrhizal Fungi.” New Phytologist 235 (6): 2158–2175. https://doi.org/10.1111/nph.18308
Cahanovitc, R., S. Livne-Luzon, R. Angel, and T. Klein. 2022. “Ectomycorrhizal fungi mediate belowground carbon transfer between pines and oaks.” The ISME Journal 16 (5): 1420–1429. https://doi.org/10.1038/s41396-022-01193-z
Chen, S., H. Zhao, C. Zou, et al. 2017. “Combined inoculation with multiple arbuscular mycorrhizal fungi improves growth, nutrient uptake and photosynthesis in cucumber seedlings.” Frontiers in Microbiology 8: 2516. https://doi.org/10.3389/fmicb.2017.02516
Chot, E., and M. S. Reddy. 2022. “Role of Ectomycorrhizal Symbiosis Behind the Host Plants Ameliorated Tolerance Against Heavy Metal Stress.” Frontiers in Microbiology 13. https://doi.org/10.3389/fmicb.2022.855473
Clemmensen, K. E., A. Bahr, O. Ovaskainen, et al. 2013. “Roots and associated fungi drive long-term carbon sequestration in boreal forest.” Science 339 (6127): 1615–1618. https://doi.org/10.1126/science.1231923
Colpaert, J. V., J. H. L. Wevers, E. Krznaric, and K. Adriaensen. 2011. “How Metal-Tolerant Ecotypes of Ectomycorrhizal Fungi Protect Plants from Heavy Metal Pollution.” Annals of Forest Science 68: 17–24. https://doi.org/10.1007/s13595-010-0003-9
Corrales, A., T. W. Henkel, and M. E. Smith. 2018. “Ectomycorrhizal Associations in the Tropics—Biogeography, Diversity Patterns and Ecosystem Roles.” New Phytoloist 220 (4): 1076–1091. https://doi.org/10.1111/nph.15151
De Quesada, G., J. Xu, Y. Salmon, et al. 2024. “The Effect of Ectomycorrhizal Fungal Exposure on Nursery-Raised Pinus sylvestris Seedlings: Plant Transpiration Under Short-Term Drought, Root Morphology and Plant Biomass.” Tree Physiology 44 (4): tpae029. https://doi.org/10.1093/treephys/tpae029
Figueiredo, A. F., J. Boy, and G. Guggenberger. 2021. “Common Mycorrhizae Network: A Review of the Theories and Mechanisms Behind Underground Interactions.” Frontiers in Fungal Biology 2: 735299. https://doi.org/10.3389/ffunb.2021.735299
Franco, A. R., N. R. Sousa, M. A. Ramos, R. S. Oliveira, and P. M. L. Castro. 2014. “Diversity and Persistence of Ectomycorrhizal Fungi and Their Effect on Nursery-Inoculated Pinus pinaster in a Post-Fire Plantation in Northern Portugal.” Microbial Ecology 68: 761–772. https://doi.org/10.1007/s00248-014-0447-9
Gehring, C. A., C. M. Sthultz, L. Flores-Rentería, A. V. Whipple, and T. G. Whitham. 2017. “Tree genetics defines fungal partner communities that may confer drought tolerance.” Proceedings National Academy of Science 114 (42): 11169–11174. https://doi.org/10.1073/pnas.1704022114
Gil-Martinez, M., Á. López-Garcia, M. T. Domínguez, et al. 2018. “Ectomycorrhizal fungal communities and their functional traits mediate plant-soil interactions in trace element contaminated soils.” Frontiers in Plant Science 9. https://doi.org/10.3389/fpls.2018.01682
Gille, C. E., P. M. Finnegan, P. E. Hayes, et al. 2023. “Facilitative and Competitive Interactions Between Mycorrhizal and Nonmycorrhizal Plants in an Extremely Phosphorus-Impoverished Environment: Role of Ectomycorrhizal Fungi and Native Oomycete Pathogens in Shaping Species Coexistence.” New Phytologist 242 (4): 1630–1644. https://doi.org/10.1111/nph.19489
Hage-Ahmed, K., K. Rosner, and S. Steinkellner. 2018. “Arbuscular Mycorrhizal Fungi and Their Response to Pesticides.” Pest Management Science 75 (3): 583–590. https://doi.org/10.1002/ps.5220
Hawkins, H.-J., R. I. M. Cargill, M. E. Van Nuland, et al. 2023. “Mycorrhizal Mycelium as a Global Carbon Pool.” Current Biology 33 (11): R560–R573. https://doi.org/10.1016/j.cub.2023.02.027
Herman, D. J., M. K. Firestone, E. Nuccio, and A. Hodge. 2011. “Interactions Between an Arbuscular Mycorrhizal Fungus and a Soil Microbial Community Mediating Litter Decomposition.” Federation of European Microbiological Societies 80 (1): 236–247. https://doi.org/10.1111/j.1574-6941.2011.01292.x
Hobbie, E. A. 2006. “Carbon allocation to ectomycorrhizal fungi correlates with belowground allocation in culture studies.” Ecology 87 (3): 563–569. https://doi.org/10.1890/05-0755
Hock, B. 2012. Fungal Associations, edited by Karl Esser. 2nd ed. Vol. 9. Springer Science & Business Media. https://doi.org/10.1007/978-3-642-30826-0
Huang, Y., X. Zhao, and S. Luan. 2007. “Uptake and Biodegradation of DDT by 4 Ectomycorrhizal Fungi.” Science of the Total Environment 385 (1–3): 235–241. https://doi.org/10.1016/j.scitotenv.2007.04.023
Godbold, D. L., G. M. Berntson, and F. A. Bazzaz. 1997. “Growth and Mycorrhizal Colonization of Three North American Tree Species Under Elevated Atmospheric CO2.” New Phytologist 137 (3): 433–440. https://doi.org/10.1046/j.1469-8137.1997.00842.x
Johnson, D., F. Martin, J. W. G. Cairney, and I. C. Anderson. 2012. “The Importance of Individuals: Intraspecific Diversity of Mycorrhizal Plants and Fungi in Ecosystems.” New Phytologist 194 (3): 614–628. https://doi.org/10.1111/j.1469-8137.2012.04087.x
Karlsen-Ayala, E., M. E. Smith, B. C. Askey, and R. Gazis. 2022. “Native ectomycorrhizal fungi from the endangered pine rocklands are superior symbionts to commercial inoculum for slash pine seedlings.” Mycorrhiza 32: 465–480. https://doi.org/10.1007/s00572-022-01092-3
Kebert, M., S. Kostić, E. Čapelja, et al. 2022. “Ectomycorrhizal Fungi Modulate Pedunculate Oak’s Heat Stress Responses through the Alternation of Polyamines, Phenolics, and Osmotica Content.” Plants 11 (23): 3360. https://doi.org/10.3390/plants11233360
Kumar, J., and N. S. Atri. 2017. “Studies on Ectomycorrhiza: An Appraisal.” Botanical Review 84: 108–155. https://doi.org/10.1007/s12229-017-9196-z
Lapeyrie, F., C. Picatto, J. Gerard, and J. Dexheimer. 1990. “T.E.M. Study of Intracellular and Extracellular Calcium Oxalate Accumulation by Ectomycorrhizal Fungi in Pure Culture or in Association with Eucalyptus Seedlings.” Symbiosis 9: 163–166.
Li, Y., T. Zhang, Y. Zhou, et al. 2021. “Ectomycorrhizal symbioses increase soil calcium availability and water use efficiency of Quercus acutissima seedlings under drought stress.” European Journal of Forest Research 140: 1039–1048.
https://doi.org/10.1007/s10342-021-01383-y
Luo, Z.-B., K. Li, Y. Gai, et al. 2011. “The ectomycorrhizal fungus (Paxillus involutus) modulates leaf physiology of poplar towards improved salt tolerance.” Environmental and Experimental Botany 72 (2): 304–311. https://doi.org/10.1016/j.envexpbot.2011.04.008
Luo, Z.-B., C. Wu, C. Zhang, H. Li, U. Lipke, and A. Polle. 2014. “The Role of Ectomycorrhizas in Heavy Metal Stress Tolerance of Host Plants.” Environmental and Experimental Botany 108: 47–62. https://doi.org/10.1016/j.envexpbot.2013.10.018
Ma, Y., J. He, C. Ma, et al. 2014. “Ectomycorrhizas with Paxillus involutus enhance cadmium uptake and tolerance in Populus × canescens.” Plant, Cell & Environment 37 (3): 627–642. https://doi.org/10.1111/pce.12183
Martin, F., A. Kohler, C. Murat, C. Veneault-Fourrey, and D. S. Hibbett. 2016. “Unearthing the Roots of Ectomycorrhizal Symbioses.” Nature Reviews Microbiology 14: 760–773. https://doi.org/10.1038/nrmicro.2016.149
Mateus, P., F. Sousa, M. Martins, et al. 2024. “The ectomycorrhizal fungus Paxillus involutus positively modulates Castanea sativa Miller (var. Marsol) responses to heat and drought co-exposure.” Plant Physiology and Biochemistry 215: 108999. https://doi.org/10.1016/j.plaphy.2024.108999
Meeds, J. A., J. M. Kranabetter, I. Zigg, et al. 2021. “Phosphorus deficiencies invoke optimal allocation of exoenzymes by ectomycorrhizas.” The ISME Journal 15: 1478–1489. https://doi.org/10.1038/s41396-020-00864-z
Nunes, L. J. R. 2023. “The Rising Threat of Atmospheric CO2: A Review on the Causes, Impacts, and Mitigation Strategies.” Environments 10 (4): 66. https://doi.org/10.3390/environments10040066
Nygren, C. M. R., J. Edqvist, M. Elfstrand, G. Heller, and A. F. S. Taylor. 2007. “Detection of Extracellular Protease Activity in Different Species and Genera of Ectomycorrhizal Fungi.” Mycorrhiza 17: 241–248. https://doi.org/10.1007/s00572-006-0100-7
Phillips, L. A., V. Ward, and M. D. Jones. 2013. “Ectomycorrhizal fungi contribute to soil organic matter cycling in sub-boreal forests.” The ISME Journal 8 (3): 699–713
https://doi.org/10.1038/ismej.2013.195
Policelli, N., T. R. Horton, A. T. Hudon, T. R. Patterson, and J. M. Bhatnagar. 2020. “Back to Roots: The Role of Ectomycorrhizal Fungi in Boreal and Temperate Forest Restoration.” Frontiers in Forests and Global Change 3. https://doi.org/10.3389/ffgc.2020.00097
Querejeta, J. I., K. Schlaeppi, Á. López-Garcia, et al. 2021. “Lower relative abundance of ectomycorrhizal fungi under a warmer and drier climate is linked to enhanced soil organic matter decomposition.” New Phytologist 232 (3): 1399–1413. https://doi.org/10.1111/nph.17661.
Ruytinx, J., L. Coninx, H. Nguyen, et al. 2017. “Identification, Evolution and Functional Characterization of Two Zn CDF‐Family Transporters of the Ectomycorrhizal Fungus Suillus luteus.” Environmental Microbiology Reports 9 (4): 419–427. https://doi.org/10.1111/1758-2229.12551
Sebastiana, M., A. B. da Silva, A. R. Matos, A. Alcântara, S. Silvestre, and R. Malhó. 2018. “Ectomycorrhizal inoculation with Pisolithus tinctorius reduces stress induced by drought in cork oak.” Mycorrhiza 28: 247–258. https://doi.org/10.1007/s00572-018-0823-2
Sevanto, S., C. A. Gehring, M. G. Ryan, et al. 2023. “Benefits of symbiotic ectomycorrhizal fungi to plant water relations depend on plant genotype in pinyon pine.” Scientific Reports 13: 14424. https://doi.org/10.1038/s41598-023-41191-5
Smith, S. E., and D. J. Read. 2008. Mycorrhizal Symbiosis. 3rd Edition. Academic Press.
Steidinger, B. S., T. W. Crowther, J. Liang, et al. 2019. “Climatic controls of decomposition drive the global biogeography of forest-tree symbioses.” Nature 569: 404–408. https://doi.org/10.1038/s41586-019-1128-0
Stuart, E. K., and K. L. Plett. 2020. “Digging Deeper: In Search of the Mechanisms of Carbon and Nitrogen Exchange in Ectomycorrhizal Symbioses.” Frontiers in Plant Science 10: 1658. https://doi.org/10.3389/fpls.2019.01658
Sun, W., B. Yang, Y. Zhu, H. Wang, G. Qin, and H. Yang. 2022. “Ectomycorrhizal fungi enhance the tolerance of phytotoxicity and cadmium accumulation in oak (Quercus acutissima Carruth.) seedlings: Modulation of growth properties and the antioxidant defense responses.” Environmental Science and Pollution Research 29: 6526–6537. https://doi.org/10.1007/s11356-021-16169-3
Tang, Y., L. Shi, K. Zhong, Z. Shen, Y. Chen. 2019. “Ectomycorrhizal fungi may not act as a barrier inhibiting host plant absorption of heavy metals.” Chemosphere 215: 115–123. https://doi.org/10.1016/j.chemosphere.2018.09.143
Thomson, B. D., T. S. Grove, N. Malajczuk, and G. E. St J. Hardy. 1994. “The Effectiveness of Ectomycorrhizal Fungi in Increasing the Growth of Eucalyptus globulus Labill. in Relation to Root Colonization and Hyphal Development in Soil.” New Phytologist 126: 517–524. https://doi.org/10.1111/j.1469-8137.1994.tb04250.x
Treseder, K. K., and S. R. Holden. 2013. “Fungal Carbon Sequestration.” Science 339 (6127): 1528–1529.
van der Heijden, M. G. A., and T. R. Horton. 2009. “Socialism in Soil? The Importance of Mycorrhizal Fungal Networks for Facilitation in Natural Ecosystems.” Journal of Ecology 97 (6): 1139–1150. https://doi.org/10.1111/j.1365-2745.2009.01570.x
van der Heijden, M. G. A., F. M. Martin, M.-A. Selosse, and I. R. Sanders. 2015. “Mycorrhizal Ecology and Evolution: The Past, the Present, and the Future.” New Phytologist 205 (4): 1406–1423. https://doi.org/10.1111/nph.13288
Vincent, B., and S. Declerck. 2021. “Ectomycorrhizal Fungi and Trees: Brothers in Arms in the Face of Anthropogenic Activities and Their Consequences.” Symbiosis 84: 337–351. https://doi.org/10.1007/s13199-021-00792-2
Vishwanathan, K., K. Zienkiewicz, Y. Liu, et al. 2020. “Ectomycorrhizal fungi induce systemic resistance against insects on a nonmycorrhizal plant in a CERK1-dependent manner.” New Phytologist 228 (2): 728–740. https://doi.org/10.1111/nph.16715
Walker, R. F. 1999. “Reforestation of an Eastern Sierra Nevada Surface Mine with Containerized Jeffrey Pine: Seedling Growth and Nutritional Responses to Controlled Release Fertilization and Ectomycorrhizal Inoculation.” Journal of Sustainable Forestry 9 (3–4): 127–147. https://doi.org/10.1300/J091v09n03_06
Wang, X. 2007. “Effects of Species Richness and Elevated Carbon Dioxide on Biomass Accumulation: A Synthesis Using Meta-Analysis.” Oecologia 152: 595–605. https://doi.org/10.1007/s00442-007-0691-5
Wang, J., H. Zhang, J. Gao, Y. Zhang, Y. Liu, and M. Tang. 2021. “Effects of Ectomycorrhizal Fungi (Suillus variegatus) on the Growth, Hydraulic Function, and Non-Structural Carbohydrates of Pinus tabulaeformis Under Drought Stress.” BMC Plant Biology 21: 171. https://doi.org/10.1186/s12870-021-02945-3
Wang, H., K. Zhang, R. Tappero, et al. 2025. “Inorganic nitrogen and organic matter jointly regulate ectomycorrhizal fungi‐mediated iron acquisition.” New Phytologist 245 (6): 2715–2725. https://doi.org/10.1111/nph.20394
Yin, D., R. Song, J. Qi, and X. Deng. 2018. “Ectomycorrhizal fungus enhances drought tolerance of Pinus sylvestris var. mongolica seedlings and improves soil condition.” Journal of Forestry Research 29: 1775–1788. https://doi.org/10.1007/s11676-017-0583-4
Yin, D., H. Wang, and J. Qi. 2021. “The Enhancement Effect of Calcium Ions on Ectomycorrhizal Fungi-Mediated Drought Resistance in Pinus sylvestris var. mongolica. Journal of Plant Growth Regulation 40: 1389–1399. https://doi.org/10.1007/s00344-020-10197-y
Zalesny, R. S., W. L. Headlee, G. Gopalakrishnan, et al. 2019. “Ecosystem Services of Poplar at Long-Term Phytoremediation Sites in the Midwest and Southeast, United States.” WIREs Energy and Environment 8 (6): e349. https://doi.org/10.1002/wene.349
Zhang, L., W. A. N. G. Ming-Xia, L. I. Hua, Y. U. A. N. Ling, J. G. Huang, and C. Penfold. 2014. “Mobilization of Inorganic Phosphorus from Soils by Ectomycorrhizal Fungi.” Pedosphere 24 (5): 683–689. https://doi.org/10.1016/S1002-0160(14)60054-0
Zhang, K., R. Tappero, J. Ruytinx, S. Branco, and H.-L. Liao. 2021. “Disentangling the Role of Ectomycorrhizal Fungi in Plant Nutrient Acquisition Along a Zn Gradient Using X-ray Imaging.” The Science of the Total Environment 801: 149481. https://doi.org/10.1016/j.scitotenv.2021.149481
Zhang, K., H. Wang, R. Tappero, et al. 2023. “Ectomycorrhizal fungi enhance pine growth by stimulating iron‐dependent mechanisms with trade‐offs in symbiotic performance.” New Phytologist 242 (4): 1645–1660. https://doi.org/10.1111/nph.19449
Zhang, W., L. Yu, B. Han, K. Liu, and X. Shao. 2022. “Mycorrhizal inoculation enhances nutrient absorption and induces insect-resistant defense of Elymus nutans.” Frontiers in Plant Science 13: 898969. https://doi.org/10.3389/fpls.2022.898969
Zheng, W., E. K. Morris, and M.C. Rillig. 2014. “Ectomycorrhizal fungi in association with Pinus sylvestris seedlings promote soil aggregation and soil water repellency.” Soil Biology and Biochemistry 78: 326–331. https://doi.org/10.1016/j.soilbio.2014.07.015
Zhu, X. C., F. B. Song, S. Q. Liu, and T. D. Liu. 2011. “Effects of Arbuscular Mycorrhizal Fungus on Photosynthesis and Water Status of Maize Under High Temperature Stress.” Plant and Soil 346: 189–199. https://doi.org/10.1007/s11104-011-0809-8

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