Open research questions in Mycorrhizal Fungi and Plant Interactions

What the literature leaves open

• Future research on medicinal plant–arbuscular mycorrhizal fungi (AMF) interactions should focus on integrating advanced molecular, ecological, and digital technologies to improve sustainable medicinal plant production and ecosystem management. High-throughput approaches such as DNA metabarcoding, whole-genome sequencing, metagenomics, transcriptomics, functional genomics and metabolomics will provide deeper insights into AMF diversity, functional traits, and plant–microbe interactions. In addition, the application of artificial intelligence (AI), remote sensing, and Geographic Information Systems (GIS) will facilitate biodiversity mapping, predictive modeling, and monitoring of ecosystem changes under climate change scenarios. Future efforts should also prioritize the indigenous AMF-based biofertilizers, development of climate-resilient carbon sequestration studies, and ecosystem restoration using native AMF communities. Translating these scientific advances into practical agricultural technologies will require strong interdisciplinary collaboration among microbiologists, botanists, ecologists, soil scientists, pharmacologists, bioinformaticians, and conservation biologists. Such integrated approaches will enhance the conservation of medicinal plant biodiversity, improve phytochemical productivity, and promote sustainable agriculture in semi-arid regions such as Marathwada (van der Heijden et al., 2015; Tedersoo et al., 2020; FAO, 2022; IPCC, 2023).

  Medicinal Plant-Arbuscular Mycorrhizal Networks in the Semi-Arid Landscapes of Marathwada: Challenges, Opportunities and Future Research · 2026 · DOI

• priorities involving molecular identification of indigenous AMF, microbiome analysis, ecological network modeling, GIS-based biodiversity mapping, climate resilience studies, region-specific mycorrhizal and biofertilizers. By from regional studies Marathwada with global advances in mycorrhizal ecology, this review provides a scientific framework for conserving medicinal plant diversity and promoting sustainable medicinal plant production under changing environmental conditions. integrating of Semi-Arid Landscapes of Marathwada The Marathwada region of Maharashtra comprises eight districts—Chhatrapati (Aurangabad), Jalna, Beed, Dharashiv (Osmanabad), Latur, Nanded, Parbhani, and Hingoli.

  Medicinal Plant-Arbuscular Mycorrhizal Networks in the Semi-Arid Landscapes of Marathwada: Challenges, Opportunities and Future Research · 2026 · DOI

• To date, the mechanism of the effect of arbuscular mycorrhizal fungus (AMF) on the Cd uptake, transport and detoxification in Solanum melongena (eggplant) remain unclear.

  Transcriptome and Physiological Analyses Show that Arbuscular Mycorrhizal Fungus Promoted Growth and Reduced Cadmium Accumulation and Phytotoxification in Solanum melongena · 2026 · DOI

• This is espe- cially fundamental if we need to access the macrofungal diversity since, compared to the extensive and long-lasting studies about microfungi diversity based on typical isola- tion techniques, there are very few studies on the diversity of macrofungi occurring in mangroves (Ghate and Sridhar 2016), and even fewer considering a global-level investi- gation (Baltazar et al.

  Worldwide diversity and ecology of mangrove fungi: a systematic review of ITS metabarcoding studies and a quantitative, integrative analysis of raw sequence data · 2026 · DOI

• Atmospheric inorganic nitrogen (N) deposition has been linked to increased tree phosphorus (P) deficiency and shifts in ectomycorrhizal (ECM) fungal community composition across Europe, but the underlying mechanisms remain poorly understood due to the scarcity of species-level studies of fungal physiology at large spatial scales.

  High nitrogen deposition is associated with phosphorus-efficient ectomycorrhizas in Europe's Scots pine forests · 2026 · DOI

• Although mycorrhizal symbiosis has been extensively studied, the complex interactive network between these fungi and MHB—which act as functional “enhancers” and “stabilizers”—and its systemic application potential remains insufficiently integrated and elucidated.

  Interactions Between Mycorrhizal Fungi and Mycorrhiza Helper Bacteria: Mechanisms, Ecological Functions, and Potential Applications in Sustainable Agriculture and Ecological Restoration · 2026 · DOI

• This review identifies priorities for the fungal community, including: (1) coordinated efforts to close major taxonomic gaps across the fungal tree of life; (2) improved repository metrics to facilitate identification of high-quality assemblies; and (3) improved and standardised genome annotation which is lacking for most assemblies.

  The impact of long-read sequencing on fungal genome assemblies: progress and disparity · 2026 · DOI

• Open Access This article is licensed under a Creative Commons Attribution-NonCommercial-NoDerivatives 4.0 International License, which permits any non-commercial use, sharing, distribution and reproduction in any medium or format, as long as you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons licence, and indicate if you modified the licensed material. You do not have permission under this licence to share adapted material derived from this article or parts of it. The images or other third party material in this article are included in the article's Creative Commons licence, unless indicated otherwise in a credit line to the material. If material is not included in the article's Creative Commons licence and your intended use is not permitted by statutory regulation or exceeds the permitted use, you will need to obtain permission directly from the copyright holder. To view a copy of this licence, visit http://creativecommons.org/licenses/by- nc-nd/4.0/. Publisher’s note Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations. © The Author(s) 2026 Communications Earth & Environment | (2026) 7:436 14

  Interactions between silicate weathering and ectomycorrhiza in severely acidified forests · 2026 · DOI

• 187. Rosling, A., Lindahl, B. D., Taylor, A. F. & Finlay, R. D. Mycelial growth and substrate acidification of ectomycorrhizal fungi in response to different minerals. FEMS Microbiol Ecol. 47, 31–37 (2004). 188. Rosling, A., Lindahl, B. D. & Finlay, R. D. Carbon allocation to ectomycorrhizal roots and mycelium colonising different mineral substrates. N. Phytol. 162, 795–802 (2004). 210. Zhu, J. et al. Changing patterns of global nitrogen deposition driven by socio-economic development. Nat. Commun. 16, 46 (2025). 211. Schulte-Uebbing, L. F., Beusen, A. H. W., Bouwman, A. F. & de Vries, W. From planetary to regional boundaries for agricultural nitrogen pollution. Nature 610, 507–512 (2022). 212. Weldon, J., Merder, J., Ferretti, M. & Grandin, U. Nitrogen deposition 189. May, H., Acker, J., Smyth, J., Bricker, O. & Dyar, M. Aqueous dissolution of low-iron chlorite in dilute acid solutions at 25 C. Clay Miner. Soc. Proc., Abstr. 32, 88 (1995). 190. Gazzè, S. A. et al. Nanoscale channels on ectomycorrhizal-colonized chlorite: Evidence for plant-driven fungal dissolution. J. Geophys. Res.: Biogeosci. 117, https://doi.org/10.1029/2012jg002016 (2012). 191. Steidinger, B. S. et al. Climatic controls of decomposition drive the global biogeography of forest-tree symbioses. Nature 569, 404–408 (2019). 192. Smith, S. E. & Read, D. J. Mycorrhizal Symbiosis. 2 edn, (1997). 193. Johnson, N. C., Gehring, C. & Jansa, J. Mycorrhizal mediation of soil: fertility, structure, and carbon storage. (Elsevier, 2016). 194. Wang, B. & Qiu, Y. L. Phylogenetic distribution and evolution of mycorrhizas in land plants. Mycorrhiza 16, 299–363 (2006). 195. Brundrett, M. C. & Tedersoo, L. Evolutionary history of mycorrhizal symbioses and global host plant diversity. N. Phytol. 220, 1108–1115 (2018). 196. Weber, G. & Claus, M. The influence of chemical soil factors on the development of VA mycorrhizas of ash (Fraxinus excelsior L.) and sycamore(Acer pseudoplatanus L.) in pot experiments. J. Plant Nutr. Soil Sci. 163, 609–616 (2000). 197. Hasselquist, N. J., Metcalfe, D. B. & Högberg, P. Contrasting effects of low and high nitrogen additions on soil CO2 flux components and ectomycorrhizal fungal sporocarp production in a boreal forest. Glob. Change Biol. 18, 3596–3605 (2012). 198. van Galen, L. G. et al. The biogeography and conservation of Earth’s ‘dark’ ectomycorrhizal fungi. Curr. Biol. 35, R563–R574 (2025). 199. Smith, S. E. & Read, D. J. Mycorrhizal symbiosis. (Academic press, 2010). 200. Martin, F., Kohler, A., Murat, C., Veneault-Fourrey, C. & Hibbett, D. S. Unearthing the roots of ectomycorrhizal symbioses. Nat. Rev. Microbiol 14, 760–773 (2016). 201. Fernandez, C. W., See, C. R. & Kennedy, P. G. Decelerated carbon cycling by ectomycorrhizal fungi is controlled by substrate quality and community composition. N. Phytol. 226, 569–582 (2020). 202. Verbruggen, E., Pena, R., Fernandez, C. W. & Soong, J. L. in Mycorrhizal Mediation of Soil 441-460 (2017). 203.

  Interactions between silicate weathering and ectomycorrhiza in severely acidified forests · 2026 · DOI

• van Diggelen, R. et al. Steenmeel en natuurherstel een gelukkige 167. Haque, F., Santos, R. M. & Chiang, Y. W. Optimizing Inorganic relatie of een risicovolle combinatie, (2019). 148. Guo, M. et al. Incorporating enhanced rock weathering into sustainable forest management. J. Environ. Manag. 394, 127335 (2025). 149. Weijters, M. et al. Herstel van droge- en vochtige heide door middel van silicaatmineralen (steenmeel). (OBN natuurkennis, 2023). 150. Van Der Bauwhede, R. et al. Restoration rocks: The long-term impact of rock dust application on soil, tree foliar nutrition, tree radial growth, and understory biodiversity in Norway spruce forest stands. Forest Ecol. Manag. 568, https://doi.org/10.1016/j.foreco.2024. 122109 (2024). 151. Van Der Bauwhede, R. et al. Three decades of monitoring the responses to alkaline K-rich applications in an acidified, K-deficient Norway spruce (Picea abies) stand reveals that forest restoration depends on treatment type: Rock dust, wood ash, dolomite and/or potassium sulphate. Geoderma 461, https://doi.org/10.1016/j. geoderma.2025.117485 (2025). 152. Lindahl, B. D. et al. Spatial separation of litter decomposition and mycorrhizal nitrogen uptake in a boreal forest. N. Phytol. 173, 611–620 (2007). 153. Van Der Bauwhede, R., van den Berg, L., Vancampenhout, K., Smolders, E. & Muys, B. Field phytometers and lab tests demonstrate that rock dust can outperform dolomite and fertilisers for acid forest soil restoration. Plant Soil. https://doi.org/10.1007/ s11104-024-07175-8 (2025). 154. Vogels, J., Bobbink, R., Weijters, M. & Bergsma, H. Steenmeeltoepassing ten behoeve van herstel biodiversiteit in Het Nationale Park De Hoge Veluwe - Eindrapport monitoring 2015- 2019. (2020). 155. Rombouts, T. (Illustrae, 2026). 156. Mikkelson, K. M., Brouillard, B. M., Bokman, C. M. & Sharp, J. O. Ecosystem Resilience and Limitations Revealed by Soil Bacterial Community Dynamics in a Bark Beetle-Impacted Forest. mBio. 8, https://doi.org/10.1128/mBio.01305-17 (2017). Carbon Sequestration and Crop Yield With Wollastonite Soil Amendment in a Microplot Study. Front Plant Sci. 11, 1012 (2020). 168. Vienne, A. et al. Earthworms in an enhanced weathering mesocosm experiment: Effects on soil carbon sequestration, base cation exchange and soil CO2 efflux. Soil Biol. Biochem. 199, https://doi. org/10.1016/j.soilbio.2024.109596 (2024). 169. Sokol, N. W. et al. Reduced accrual of mineral-associated organic matter after two years of enhanced rock weathering in cropland soils, though no net losses of soil organic carbon. Biogeochemistry 167, 989–1005 (2024). 170. Zhu, E. et al. Enhanced Mineral Preservation Rather Than Microbial Residue Production Dictates the Accrual of Mineral-Associated Organic Carbon Along a Weathering Gradient. Geophys. Res. Lett. 51, https://doi.org/10.1029/2024gl108466 (2024). 171. Liang, G., Stark, J. & Waring, B. G. Mineral reactivity determines root effects on soil organic carbon. Nat. Commun.

  Interactions between silicate weathering and ectomycorrhiza in severely acidified forests · 2026 · DOI

• van Breemen, N. et al. Mycorrhizal weathering: A true case of mineral 125. Kjøller, R. & Clemmensen, K. E. Belowground ectomycorrhizal plant nutrition? Biochemistry 49, 53–67 (2000). 105. Kinraide, T. B. Toxicity factors in acidic forest soils: attempts to fungal communities respond to liming in three southern Swedish coniferous forest stands. For. Ecol. Manag. 257, 2217–2225 (2009). evaluate separately the toxic effects of excessive Al3+ and H+ and insufficient Ca2+ and Mg2+ upon root elongation. Eur. J. Soil Sci. 54, 323–333 (2008). 126. Pabian, S. E., Rummel, S. M., Sharpe, W. E. & Brittingham, M. C. Terrestrial Liming As a Restoration Technique for Acidified Forest Ecosystems. Int. J. Forestry Res. 2012, 1–10 (2012). 106. Cumming, J. R. & Weinstein, L. H. Aluminum-mycorrhizal interactions in the physiology of pitch pine seedlings. Plant Soil 125 (1990). 107. Cumming, J. R. & Weinstein, L. H. Utilization of AlPO4 as a phosphorus source by ectomycorrhizal Pinus rigida Mill. seedlings. N. Phytologist 116, 99–106 (1990). 108. Göransson, A. & Eldhuset, T. D. Is the Ca+K+Mg/Al ratio in the soil solution a predictive tool for estimating forest damage? Water, Air, Soil Poll. 1 (2001). 109. Hentschel, E., Godbold, D. L., Marschner, P., Schlegel, H. & Jentschke, G. The effect of Paxillus involutus Fr. on aluminum sensitivity of Norway spruce seedlings. Tree Physiol. 12, 379–390 (1993). 110. Schier, G. A. & Mcquattie, C. J. Effect of aluminum on the growth, anatomy, and nutrient content of ectomycorrhizal and nonmycorrhizal eastern white pine seedlings. Canadian J. Forest Res. 25, 1252–1262 (1995). 111. Verbruggen, E. (BioRender, 2026). 112. Reid, C. & Watmough, S. A. Evaluating the effects of liming and wood-ash treatment on forest ecosystems through systematic meta-analysis. Can. J. For. Res. 44, 867–885 (2014). 113. Jansone, L., von Wilpert, K. & Hartmann, P. Natural Recovery and Liming Effects in Acidified Forest Soils in SW-Germany. Soil Syst. 4, https://doi.org/10.3390/soilsystems4030038 (2020). 114. Nilsson, S. I. et al. Influence of dolomite lime on leaching and storage of C, N and S in a Spodosol under Norway spruce (Picea abies (L.) Karst. For. Ecol. Manag. 146, 55–73 (2001). 127. Clivot, H. et al. Changes in soil bacterial communities following liming of acidified forests. Appl. Soil Ecol. 59, 116–123 (2012). 128. Holland, J. E. et al. Liming impacts on soils, crops and biodiversity in the UK: A review. Sci. Total Environ. 610-611, 316–332 (2018). 129. Baumann, M., Dittrich, S., Körner, M. & von Oheimb, G. Liming in spruce stands: What effect does the number of lime applications have on the herb layer? Eur. J. For. Res. 138, 723–735 (2019). 130. Greve, M., Block, J., Schüler, G. & Werner, W. Long Term Effects of Forest Liming on the Acid-Base Budget. Appl. Sci. 11, https://doi. org/10.3390/app11030955 (2021). 131. Sridhar, B. et al.

  Interactions between silicate weathering and ectomycorrhiza in severely acidified forests · 2026 · DOI

• 59. 60. 61. 62. Treseder, K. K. A meta-analysis of mycorrhizal responses to nitrogen, phosphorus, and atmospheric CO(2) in field studies. N. Phytol. 164, 347–355 (2004). Li, W. et al. The effects of simulated nitrogen deposition on plant root traits: A meta-analysis. Soil Biol. Biochem. 82, 112–118 (2015). Kowalski, S., Obloza, E. & Wojewoda, W. Susceptibility of ectomycorrhizal and ectendomycorrhizal fungi to pH of the enviroment. Acta Mycologica 31, 127–136 (1996). Jongbloed, R. H. Response of isolated ectomycorrhizal fungi to chemical stress factors of acid soils, University of Nijmegen, (1992). 64. 63. De Schrijver, A. et al. Acidification of forested podzols in North Belgium during the period 1950-2000. Sci. Total Environ. 361, 189–195 (2006). Zhang, Y., Zhou, J. & Chen, H. Global Trajectories of Forest Soil Acidification: A Scientometric Synthesis of Drivers, Impacts and Sustainable Solutions. Forests 16, https://doi.org/10.3390/ f16050733 (2025). Achilles, F. et al. Effects of Moderate Nitrate and Low Sulphate Depositions on the Status of Soil Base Cation Pools and Recent Mineral Soil Acidification at Forest Conversion Sites with European Beech (“Green Eyes”) Embedded in Norway Spruce and Scots Pine Stands. Forests 12, https://doi.org/10.3390/f12050573 (2021). 66. Hopf, S.-E. et al. Dendrochemical indicators of tree rings reveal 65. historical soil acidification in Swiss forest stands. Dendrochronologia 81, https://doi.org/10.1016/j.dendro.2023. 126099 (2023). Tian, D. & Niu, S. A global analysis of soil acidification caused by nitrogen addition. Environ. Res. Lett. 10, https://doi.org/10.1088/ 1748-9326/10/2/024019 (2015). 67. 68. Maes, J. et al. Accounting for forest condition in Europe based on an 69. 70. 71. 72. international statistical standard. Nat. Commun. 14, 3723 (2023). Amacher, M. C., O’Dea, C. & Page-Dumroese, D. Indicator 4.19: Area and percent of forest land with significant soil degradation. (2020). Van Der Bauwhede, R., Muys, B., Vancampenhout, K. & Smolders, E. Rock dust as a restoration measure for acidified forests: from mineral to helicopter. (2025). Kochian, L. Cellular mechanisms of aluminum toxicity and resistance in plants. Plant Physiol. Plant Mol. Biol. 46, 237–260 (1995). Treseder, K. K. Nitrogen additions and microbial biomass: a metaanalysis of ecosystem studies. Ecol. Lett. 11, 1111–1120 (2008). 74. 76. 73. Gundersen, P. & Rasmussen, L. Nitrification in Forest Soils: Effect from Nitrogen Deposition on Soil Acidification and Aluminum Release. Rev. Environ. Contamination Toxicol. 113, 1–2 (1990). Abrahamsen, G. Effects of acidic desposition of forest soil and vegetation. Philos. Trans. R. Soc. Lond. 305, 369–382 (1984). 75. Huettl, R. F. & Zoettl, H. W. Liming as a mitigation tool in Germany’s declining forest - reviewing results from former and recent trials. For. Ecol. Manag. 61, 325–338 (1993). Schlesinger, W. Biogeochemistry: an analysis of global change. (1997). Bergsma, H. et al.

  Interactions between silicate weathering and ectomycorrhiza in severely acidified forests · 2026 · DOI

• 16. 17. Brantley, S. L. et al. Twelve testable hypotheses on the geobiology of weathering. Geobiology 9, 140–165 (2011). Li, Q. et al. Soil acidification of the soil profile across Chengdu Plain of China from the 1980s to 2010s. Sci. Total Environ. 698, 134320 (2020). 37. 38. 18. Guo, J. H. et al. Significant acidification in major Chinese croplands. 39. Science 327, 1008–1010 (2010). 20. 19. Haselwandter, K. Structure and function of siderophores produced by mycorrhizal fungi. Mineralogical Mag. 72, 61–64 (2008). van Scholl, L., Smits, M. M. & Hoffland, E. Ectomycorrhizal weathering of the soil minerals muscovite and hornblende. N. Phytol. 171, 805–813 (2006). Phillips, J. D. Biological energy in landscape evolution. Am. J. Sci. 309, 271–289 (2009). 21. 22. Heinemeyer, A. et al. Exploring the “overflow tap” theory: linking 23. 24. forest soil CO2 fluxes and individual mycorrhizosphere components to photosynthesis. Biogeosciences 9, 79–95 (2012). Taylor, L. L., Banwart, S. A., Valdes, P. J., Leake, J. R. & Beerling, D. J. Evaluating the effects of terrestrial ecosystems, climate and carbon dioxide on weathering over geological time: a global-scale process-based approach. Philos. Trans. R. Soc. Lond. B Biol. Sci. 367, 565–582 (2012). Taylor, L., Banwart, S., Leake, J. & Beerling, D. J. Modeling the evolutionary rise of ectomycorrhiza on sub-surface weathering environments and the geochemical carbon cycle. Am. J. Sci. 311, 369–403 (2011). 26. 25. Wu, Z. et al. Soil acidification and factors controlling topsoil pH shift of cropland in central China from 2008 to 2018. Geoderma 408, https://doi.org/10.1016/j.geoderma.2021.115586 (2022). Landeweert, R., Hoffland, E., Finlay, R., Kuyper, T. W. & Breemen, N. v. Linking plants to rocks:ectomycorrhizal fungi mobilize nutrients from minerals. Trends Ecol Evol. 16, 248–254 (2001). van Schöll, L. et al. Rock-eating mycorrhizas: their role in plant nutrition and biogeochemical cycles. Plant Soil 303, 35–47 (2007). Thorley, R. M., Taylor, L. L., Banwart, S. A., Leake, J. R. & Beerling, D. J. The role of forest trees and their mycorrhizal fungi in carbonate rock weathering and its significance for global carbon cycling. Plant Cell Environ. 38, 1947–1961 (2015). 28. 27. 29. Hoffland, E., Giesler, R., Breemen, N. V. & Jongmans, A. G. Feldspar Tunneling by Fungi along Natural Productivity Gradients. Ecosystems 6, 739–746 (2003). 33. 31. 32. 30. Wallander, H. Uptake of P from apatite by Pinus sylvestris seedlings colonised by different ectomycorrhizal fungi. Plant Soil 218, 249–256 (1999). Rosenstock, N. P., Berner, C., Smits, M. M., Kram, P. & Wallander, H. The role of phosphorus, magnesium and potassium availability in soil fungal exploration of mineral nutrient sources in Norway spruce forests. N. Phytol. 211, 542–553 (2016). Smits, M. M. & Wallander, H. in Mycorrhizal Mediation of Soil 35–46 (2017). Smits, M. M., Johansson, L. & Wallander, H.

  Interactions between silicate weathering and ectomycorrhiza in severely acidified forests · 2026 · DOI

• Upscaling and choice of minerals. Scaling up from plot-level experi- ments to stand- and landscape-level assessments will be crucial. Only by integrating diverse forest types, soil chemistries, and ESW amendments can we determine the generality of effects and assess the true potential of basalt–mycorrhiza interactions in acidified environments. As shown in Table 1, mineral choice is critical. Some minerals, such as quartz, dissolve too slowly and lack the basic cations that EcM fungi can exploit, whereas others, including calcite and dolomite, dissolve too rapidly, potentially leading to ecological imbalances. Minerals containing nutrients of interest for EcM fungi (e.g., Mg, K, P) tend to show higher weathering activities than those without165. Minerals of particular interest include leucite, anorthite, phlogopite, biotite, and muscovite, which have been shown to be actively weathered by EcM and possess substantial buffering potential. Beyond mineral choice, it is important to consider that in extremely acidified systems with high N deposition rates there can be a large organic N pool of undecomposed litter. When pH increases, organic matter decom- position will be stimulated, releasing substantial amounts of N that remain available for extended periods; upon nitrification and subsequent leaching from the system, this N can induce additional acidification. How long this increased inorganic N concentration will interfere with EcM fungal restoration is an urgent question. Moreover, current N deposition will certainly lengthen this period, if not interfere with recovery indefinitely. Hence, it will be important to assess whether the subset of “nitrophilic” EcM taxa96 that remains in the ESW- treated system is sufficiently capable of stimulating mineral weathering, or whether the increased N availability will have an overall negative effect on weathering potential. Furthermore, whether recovery timelines can be shortened by assisted migration (e.g., inoculation) of EcM fungal species that have disappeared due to chronic N deposition is an urgent subject of future studies. ESW–EcM interactions in soil carbon cycling. Lastly, an important research opportunity lies in establishing how EcM modulates the potential effect of ESW on soil carbon cycling. Recent ESW experiments (with wollastonite) indicate that mineral nutrient release (particularly Ca, Si, and Fe) can promote SOC sequestration in stable MAOM fractions more effectively than it promotes inorganic C capture166. In strongly acidified forest soils, where weathering may partly proceed via strong- acid pathways and where released base cations can be temporarily retained on soil exchange complexes, the inorganic CDR potential of ESW may be reduced or delayed. However, these conditions do not preclude significant ecosystem benefits, including improvements in soil buffering capacity, nutrient availability, and potential organic carbon stabilisation. EcM fungi may also further enhance SOM build-up and stabilisation through extracellular enzyme exudation and organic N acquisition pathways. In addition, mineral nutrients released from cru- shed rocks can stimulate plant growth140,167, thereby enhancing litter production causing long-term increases in soil C stock (after the recovery period, see above), as well as altering organic matter transformations by soil biota168. These processes promote MAOM formation either directly through the binding of plant- and microbial-derived residues169 to mineral elements170,171, or indirectly via aggregation with secondary minerals formed during weathering170,172,173. However, in-field evidence for these processes following ESW remains scarce, and the role of EcM fungi within them is still unclear. This gap needs to be urgently filled to discover possible synergies between ESW mitigation of acidification and its C sequestration potential.

  Interactions between silicate weathering and ectomycorrhiza in severely acidified forests · 2026 · DOI

• Fig. 3 | Conceptual schematic of the hypothesised effects of silicate rock dust application on acidic, poorly buffered forest soils. Silicate amendment is expected to raise soil pH (pH-0.01 M CaCl₂ > 3.5), increase base cations, reduce toxic Al³⁺ availability, and promote ectomycorrhizal (EcM) colonisation compared to untreated acidic soils (pH-0.01 M < 3.3), where base cations are depleted and Al³⁺ dominates. Insets illustrate (a) untreated, poorly buffered soils with a small cation exchange complex (CEC) and a low base cation/Al³⁺ ratio, and (b) silicate-amended soils with higher base cation / Al³⁺ ratio. Plant roots are shown in brown. Green circles represent base cations (Ca²⁺, Mg²⁺, K⁺), red circles Al³⁺, and grey circles H⁺. The ‘Cation exchange complex’ is schematically depicted and represents all reactive soil surfaces (including clay minerals and organic matter). Black rectangles indicate primary silicate minerals (e.g., K-bearing feldspars) present in base-poor soils. Grey polygons represent added silicate rock dust particles (ESW). Arrows indicate the flux of photosynthetic C from roots to EcM, the exudation of enzymes and organic acids, and the consequent weathering of primary minerals leading to base cation release. Created by the authors using Illustrae.

  Interactions between silicate weathering and ectomycorrhiza in severely acidified forests · 2026 · DOI

• EcM fungi themselves could enhance the weathering of applied silicates (see above), creating a positive feedback between fungal activity, mineral amendment, nutrient mobilisation, and overall soil health. In this way, ESW aligns naturally with EcM-mediated soil processes, potentially rebalancing the nutrients in ecosystems destabilised by chronic N-driven acidification. Current state-of-art of ESW application in forests Although rock dust applications were already known in the first half of the twentieth century in forestry and agriculture, ESW has recently gained considerable attention as a CO2 removal (CDR) strategy in agricultural contexts142–144. While its potential for long-term C sequestration is a key aspect, the original application of ESW also provides multiple benefits for ecosystem restoration. Within the forest sector of the Netherlands and Belgium, for instance, ESW has been defined as a ‘no-regret’ measure against acidification, and its application is gaining momentum in large-scale forest145–148 and heathland restoration projects149. Comparable mineral amendments have precedent in North America, at both experimental and forest management application level. For example, the watershed-scale calcium silicate addition experiment at the Hubbard Brook Experimental Forest demonstrated that silicate applications can substantially improve nutrient cycling and soil chemistry in acidified forest ecosystems141. While this experiment was designed as an ecosystem manipulation rather than a scalable deployment strategy, other studies have explored practical imple- mentation approaches. For example, Guo et al. 148 demonstrated that silicate rock powders can be integrated into existing forest management operations using harvesting trail networks and ground-based spreading equipment148. Recent field studies demonstrate that ESW can indeed improve buf- fering in acidified and N-saturated forest soils. Two long-term studies revisited Norway spruce (Picea abies L.) stands about three decades after an application of 4.7 Mg basalt/ha or 10 Mg phonolite/ha, and reported long- term significant improvements in ecosystem functioning150,151. These included increases in pH, effective CEC, elevated base cation concentrations (Ca2+, Mg2+, K+), and reduced Al saturation – especially in the upper mineral soil layers most relevant to EcM functioning152. Moreover, over time, base cations reached deeper soil horizons, and foliar nutrient imbal- ances in these conifers were partially corrected. Positive effects also com- prised improved vitality, increased wood volume production, reduced defoliation and stronger growth, particularly among younger trees and under higher N loads150. Complementary trials showed similar growth- improving patterns on planted saplings, even under closed-canopy conditions153. In addition, increases in understory plant diversity were evi- denced without the loss of calcifuge species.150. Together, these long-term observations suggest that single silicate rock dust applications can provide pH buffering effects lasting decades, implying that repeated annual appli- cations are not required in forest ecosystems, and reapplication is likely only needed after 50+ years70. Conversely, higher or repeated applications are not without risk. The response of EcM communities to sustained inputs remains uncertain, and excessive application rates may induce effects similar to liming, including altered nutrient cycling and increased N losses.

  Interactions between silicate weathering and ectomycorrhiza in severely acidified forests · 2026 · DOI

• pH 3.561,62. Such thresholds are routinely exceeded in acidified forest soils, with severely acidified sites showing topsoil pH-0.01 M CaCl2 < 3.3 on poorly buffered sandy soils, for example, in parts of Western Europe. Above their physiological limit (pH > 3.5), EcM fungi can play an important role in mitigating the adverse effects of acidification27. By releasing base cations through mineral weathering, EcM fungi contribute to long-term soil pH stabilisation104. In strongly acidic soils (pH < 4), EcM can also mitigate Al3+ toxicity by oxalate exudation, particularly when Mg or P is limiting20,104,105. EcM-colonised seedlings have higher growth rates than non-mycorrhizal seedlings under Al stress, potentially due to (i) better nutrient supply, (ii) metal detoxification, and (iii) replacement of exposed root surfaces by protective mantle-sheeted mycorrhiza27,106–110. In spite of such compensatory mechan- isms, EcM still have clear physiological constraints under low-pH conditions, which is compounded by reduced performance of their hosts reducing C allocation from trees to their fungal symbiont (Fig. 2)111. This is why any attempt to restore forest health must address both the chemical recovery of the soil and the biological restoration of these vital fungal communities. Amendments against soil acidification Because EcM fungi and trees suffer under severe acidification, it is important to increase the forest soil’s buffering capacity to more favourable conditions. Forest managers have long applied liming (typically using calcite or dolo- mite) and mineral fertilisers (KCl, K2SO4, a.o.) to raise soil pH and replenish nutrient pools112. Forest liming has a long history in Europe, originating in the early 20th century in Germany where it was used to counteract natural soil acidity and improve forest productivity. From the 1950s onwards, systematic experiments in Finland and Sweden further explored its role in enhancing tree growth and stimulating nitrogen mineralization in forest soils with intrinsic slow decomposition rate112,113. The focus shifted in the 1980s, when liming began to be applied at a wider scale to mitigate soil acidification caused by atmospheric deposition and acid rain75,113,114. Since then, it has remained an important management practice in Central and Northern Europe to restore soil buffering capacity and address nutrient imbalances in forest ecosystems75,112–121 with large-scale implementation in countries such as Germany, where approximately 29% of forest area (10.9 million hectares) had been limed by 2013122. Liming increases soil pH and BS by supplying Ca2+ (and Mg2+), which displace H+ and Al3+ from exchange sites and reduce aluminium toxicity115,123. Across long-term experiments, pH increases of roughly 0.1–2.4 units and substantial improvements in BS have been reported123,124. However, conventional approaches like liming do not supply aluminosili- cates. Moreover, their aggressive dissolution rates come with ecological costs. For example, liming can accelerate organic matter decomposition and nitrification, while promoting the dominance of ruderal species124–132. In particular, EcM communities often show drastic shifts in composition after liming, without necessarily gaining in species richness. Dolomite liming favours saprotrophic fungi and suppresses acidophilic EcM species, either due to increased N availability or a sudden pH increase125,133–135. Given these limitations, attention is turning to alternative, less dis- ruptive methods to restore buffering in acidified forest soils. One such approach is Enhanced Silicate Weathering (ESW): the application of finely ground silicate rock dust that acts as a slow-release soil amendment.

  Interactions between silicate weathering and ectomycorrhiza in severely acidified forests · 2026 · DOI

• Box 2: | Global context of soil acidification Over the past four decades, global atmospheric N deposition has increased by about 8%. N deposition is in the form of NOx and reduced N (NH₃, NH₄⁺), mainly from fossil fuel combustion, fertiliser use, and animal husbandry, with the highest levels in densely populated areas such as East Asia205. Deposition patterns vary regionally. Following peaks around 1990, NOx emissions declined sharply in Europe, North America, Korea and Japan, owing to stricter air quality controls205–209. In contrast, reduced N forms (NH₃, NH₄⁺) from intensive livestock farming and overfertilisation still exceed critical loads in various regions, for example in parts of northwestern Europe (Poland, Denmark, Germany, the Benelux)210. Present hotspots of total N exceedance in reference to the critical N surplus include India, Pakistan, and eastern China, with smaller sources in the Nile Basin, the Peruvian coast, and parts of Saudi Arabia211. As a result, many ecosystems, particularly lowland habitats, continue to surpass critical N loads, affecting plant vitality, vegetation structure, species composition, and soil chemistry212–216. Elevated N deposition remains a major cause of soil acidification217. Historically, sulphur (S) inputs (e.g., SO₂) also contributed significantly to acidification, but S deposition has declined in Europe and North America nutrient equilibria that EcM fungi need and help to sustain57–60. In particular, the buffering role of EcM-mediated weathering is being undermined by rapid base cation leaching and depletion of primary minerals and aluminium toxicity, which destabilise the fungal networks61,62. As a consequence, mycorrhiza-mediated nutrient cycling and plant nutrient acquisition may be compromised, with potential cascading effects on soil processes and plant vitality56,57,59. In the following, we focus on how N deposition and acidification affect extratropical forests dominated by EcM fungi (Box 2). At the global scale, N deposition is a major driver of soil acidification, with widespread impacts reported across regions including China, the US, and Central Europe63–67. A global meta-analysis showed that N addition reduced soil pH by an average of 0.26 pH units across terrestrial ecosystems, with similarly negative but variable responses in forest ecosystems worldwide67. For example, critical loads for acidification are exceeded across substantial forest areas, affecting an estimated 21% of forests in the United States (2020) and 30% in the European Union (2016)68,69. In forests, N deposition contributes to soil acidification primarily through nitrification, during which protons (H+) are produced as ammonium is oxidised to nitrate. When nitrate is not taken up by plants, it leaches from the root zone, causing a net loss of base cations (Ca2+, Mg2+ and K+), thereby further reducing the soil pH buffering capacity (Fig. 2).

  Interactions between silicate weathering and ectomycorrhiza in severely acidified forests · 2026 · DOI

• Box 1: | Ectomycorrhizal (EcM) fungi Mycorrhizal associations first evolved in the early Devonian Rhynie chert (~411 Ma), and today they occur between ~50,000 fungal and 250,000 plant species 6,7. Ectomycorrhizal symbiosis – one of four general mycorrhiza types – arose only in the Cretaceous and is formed between ectomycorrhizal fungi and the roots of predominantly long-lived woody perennials. Although fewer than 5% of terrestrial plant species engage in this symbiosis26, it characterises the dominant trees of temperate forests (Pinaceae, Fagaceae), several (sub)tropical families (e.g. Fabaceae, Dipterocarpaceae), and especially high-latitude boreal forests26,191,192. EcM fungi form essential partnerships with ~8,500 species of angiosperms (28 lineages) and gymnosperms (2 lineages). They are most dominant in ecosystems where cold or dry conditions limit decomposition, rendering tree growth and survival highly dependent on fungal partners193–195. In boreal forests, for example, at least 95% of root tips are colonised by EcM fungi192,196, which account for 30-40% of soil microbial biomass197. An estimated 25,000 EcM fungal species – mainly Basidiomycota and Ascomycota – form structures distinct from those of arbuscular mycorrhizal (AM) fungi, the most ancient and widespread mycorrhizal type1,26,192,198. Unlike AM fungi, EcM hyphae do not penetrate plant cells but instead form a Hartig net surrounding root cortical cells199–201 (Fig. 1). Extensive extraradical hyphae expand the plant’s depletion zone into bulk soil, accessing microsites beyond the reach of roots26. With their high surface-area-to-mass ratio, EcM fungi efficiently scavenge nutrients, while receiving 10–30% of the host’s photosynthetically fixed carbon16,26,192. Although AM fungi share this bidirectional exchange, EcM fungi – derived from multiple lineages of higher fungi – possess a far broader enzymatic capacity, allowing them to mobilise organic N and P, a trait largely absent in AM fungi35,202–204. to plant roots1,26,27. EcM fungi contribute to physical weathering by extending their fine hyphae (3–10 μm in diameter) into rock pores and fissures, where high turgor pressure enlarges pre-existing cracks and cavities7,26,28,29. Secondly, accelerated chemical weathering is often regarded as one of the most important biological weathering processes in soils26,28. EcM fungi facilitate accelerated chemical weathering by dissolving primary minerals (e.g., feldspars, plagioclases and apatite) through the exudation of low-molecular-weight organic acids (LMWOAs) like oxalate, citrate, and malate17,20,26,28,30–32. However, the actual contribution of LMWOAs to mineral dissolution remains debated: while concentrations in bulk soil are generally too low to drive measurable weathering, microscale accumulation within the rhizosphere or hyphosphere may reach levels sufficient to enhance mineral dissolution33.

  Interactions between silicate weathering and ectomycorrhiza in severely acidified forests · 2026 · DOI

• This study has certain limitations. Although a paired sampling design (cultivated soil vs. adjacent uncultivated control soil) was employed, which effectively eliminates background differences inherent to the sampling sites themselves to isolate plant-induced changes, it must be acknowledged that each medicinal plant species was sourced from its respective primary, yet geographically separate, cultivation area. Consequently, plant species and geographic environmental factors (e.g., climate, soil parent material) are confounded to some extent. While climatic factors were included in the analysis, and the three Panax species, originating from different climatic zones, exhibited similar trends—providing some support for the dominant role of plant species—the potential influence of regional environmental background on the observed interspecific differences cannot be entirely ruled out.

  Ecological strategies determine continuous cropping susceptibility in Panax and Achyranthes · 2026 · DOI

• The research focuses exclusively on rhizosphere fungal diversity in maize but does not examine how fungal community differences translate across different maize varieties, soil types, or geographic regions. Validation of whether the observed polyculture-driven fungal profiles are reproducible across diverse maize cultivars and edaphic conditions is needed for sustainable agriculture applications.

  Metagenomic Analysis of Rhizosphere Fungal Diversity in Maize with Various Crop Systems · 2026 · DOI

• The study identifies marker genes associated with microbial quantity, relative abundance, and alpha diversity in rhizosphere fungal communities but does not establish causal relationships between specific fungal taxa or gene variants and measurable plant phenotypes (yield, disease resistance, nutrient uptake). Experimental validation linking dominant fungal profiles from polyculture versus monoculture systems to plant performance metrics is absent.

  Metagenomic Analysis of Rhizosphere Fungal Diversity in Maize with Various Crop Systems · 2026 · DOI

• The paper acknowledges that many of the identified fungi cannot be cultured using conventional methods, limiting validation of their functional roles and development as biocontrol agents. Direct cultivation isolation techniques, synthetic media optimization, or co-culture with maize roots are not explored to enable laboratory characterization of the unculturable fungal taxa identified in the metagenomic analysis.

  Metagenomic Analysis of Rhizosphere Fungal Diversity in Maize with Various Crop Systems · 2026 · DOI

• The study demonstrates differences in fungal community composition between monoculture and polyculture maize systems but does not include temporal or seasonal sampling across multiple growing seasons. Long-term monitoring of these rhizosphere fungal communities under different crop rotation schedules and environmental conditions is needed to determine stability and predictability of the observed fungal profiles.

  Metagenomic Analysis of Rhizosphere Fungal Diversity in Maize with Various Crop Systems · 2026 · DOI

• The paper identifies functionally important fungi with potential roles in nutrient cycling, plant growth promotion, and pathogen suppression in maize rhizosphere communities, but does not characterize the specific metabolic pathways or functional gene annotations of these taxa. Genomic mapping is mentioned as critical, yet the study lacks functional metagenomics or whole-genome sequencing data to validate the actual biocontrol mechanisms and nutrient cycling capabilities of the dominant fungal groups identified.

  Metagenomic Analysis of Rhizosphere Fungal Diversity in Maize with Various Crop Systems · 2026 · DOI

Most-cited papers in Mycorrhizal Fungi and Plant Interactions

• Global diversity and geography of soil fungi · Science · 2014 · 3,218 citations
• Mechanisms and implications of bacterial–fungal competition for soil resources · The ISME Journal · 2024 · 553 citations
• Cómo cambiar el mundo · Harvard business review · 2008 · 256 citations
• The mycorrhizal symbiosis: research frontiers in genomics, ecology, and agricultural application · New Phytologist · 2024 · 229 citations
• The enzyme patterns of Ascomycota and Basidiomycota fungi reveal their different functions in soil · Applied Soil Ecology · 2024 · 158 citations
• Above-and below-ground feedback loop of maize is jointly enhanced by plant growth-promoting rhizobacteria and arbuscular mycorrhizal fungi in drier soil · The Science of The Total Environment · 2024 · 102 citations
• A tripartite bacterial-fungal-plant symbiosis in the mycorrhiza-shaped microbiome drives plant growth and mycorrhization · Microbiome · 2024 · 100 citations
• Current trends, limitations and future research in the fungi? · Fungal Diversity · 2024 · 94 citations
• Enhancing soil health and nutrient cycling through soil amendments: Improving the synergy of bacteria and fungi · The Science of The Total Environment · 2024 · 94 citations
• Distribution and drivers of ectomycorrhizal fungal communities across the North American Arctic · Ecosphere · 2012 · 94 citations

Most recent work

• Ecological strategies determine continuous cropping susceptibility in Panax and Achyranthes · Frontiers in Plant Science · 2026
• Metagenomic Analysis of Rhizosphere Fungal Diversity in Maize with Various Crop Systems · International Journal of Agriculture and Biosciences · 2026
• Impacts of ectomycorrhizal forest change on nutrient cycling, forest resilience, and ecosystem services · The ISME Journal · 2026
• Using pathogen nutrient acquisition strategy to explain the occurrence of fungal and oomycete diseases in wild plants · Philosophical Transactions of the Royal Society B Biological Sciences · 2026
• "Fuzzy specificity" shapes diazotroph diversity and composition in nodulating plants of the Southeastern USA · bioRxiv · 2026
• How arbuscular mycorrhizal fungi maintain plant nitrogen acquisition under drought · Biology and Fertility of Soils · 2026
• Evaluation of Innovative Cultivation Practices for Optimizing Soil Fertility, Crop Yields and Soil Microorganisms in Western Niger · International Journal of Plant & Soil Science · 2026
• Seeing the invisible: A new genus and species of Sebacinales (Basidiomycota) with invisible basidiomata · Mycological Progress · 2026
• <i>Pluteus sinensis</i> , a New Species of <i>Pluteus</i> Sect. <i>Celluloderma</i> from China · New Zealand Journal of Botany · 2026
• Arbuscular mycorrhizal symbiosis enhances the cadmium stress tolerance of Medicago sativa in association with regulation of the aminolevulinic acid pathway · BMC Plant Biology · 2026

Related topics in Agricultural and Biological Sciences