Mycorrhizal colonization of dryland tree establishment depends on soil microbial cooperation

1. Ren, S. et al. Storage potential of soil functional carbon fractions in the World’s largest plantations. Adv. Sci. 12, e04995 (2025).

   Google Scholar 

2. Wang, Y. J. et al. Land availability and policy commitments limit global climate mitigation from forestation. Science 389, 931–934 (2025).

   Google Scholar 

3. Garrett, L., Lévite, H., Besacier, C., Alekseeva, N. & Duchelle, M. The key role of forest and landscape restoration in climate action. (FAO, 2022).

4. Reynolds, J. F. et al. Global desertification: building a science for dryland development. Science 316, 847–851 (2007).

   Google Scholar 

5. Ma, L. L. et al. Planted forests in China have higher drought risk than natural forests. Glob. Change Biol. 31, e70055 (2025).

   Google Scholar 

6. Koppa, A. et al. Dryland self-expansion enabled by land-atmosphere feedbacks. Science 385, 967–972 (2024).

   Google Scholar 

7. Gu, L., Schumacher, D. L., Wang, H. M., Yin, J. & Fischer, E. M. Land-atmosphere feedbacks drive dryland drought and expansion under climate warming. Innovation 6, 100863 (2025).

   Google Scholar 

8. Cosme, M. Mycorrhizas drive the evolution of plant adaptation to drought. Commun. Biol. 6, 346 (2023).

   Google Scholar 

9. Sachsenmaier, L. et al. Forest growth resistance and resilience to the 2018–2020 drought depend on tree diversity and mycorrhizal type. J. Ecol. 112, 1787–1803 (2024).

   Google Scholar 

10. Poudel, M. et al. The role of plant-associated bacteria, fungi, and viruses in drought stress mitigation. Front. Microbiol. 12, 743512 (2021).

    Google Scholar 

11. Hu, M. Y. et al. Plant functional traits affect biomass responses to global change: a meta-analysis. J. Ecol. 113, 2046–2065 (2025).

    Google Scholar 

12. Van Nuland, M. E. et al. Global hotspots of mycorrhizal fungal richness are poorly protected. Nature 645, 414–422 (2025).

    Google Scholar 

13. Jiang, F. et al. Mycorrhizal symbioses and tree diversity in global forest communities. Sci. Adv. 11, eadt5743 (2025).

    Google Scholar 

14. Berdugo, M. et al. Global ecosystem thresholds driven by aridity. Science 367, 787–790 (2020).

    Google Scholar 

15. Stuart, E. K. et al. Acquisition of host-derived carbon in biomass of the ectomycorrhizal fungus Pisolithus microcarpus is correlated to fungal carbon demand and plant defences. FEMS Microbiol. Ecol. 99, fiad037 (2023).

    Google Scholar 

16. Canarini, A. et al. Soil fungi remain active and invest in storage compounds during drought, independent of future climate conditions. Nat. Commun. 15, 10410 (2024).

    Google Scholar 

17. Xie, L. L. et al. Variations in ectomycorrhizal exploration types parallel seedling fine root traits of two temperate tree species under extreme drought and contrasting solar radiation treatments. Plant Cell Environ. 47, 5053–5066 (2024).

    Google Scholar 

18. Guarnizo, A. L., Navarro-Ródenas, A., Calvo-Polanco, M., Marqués-Gálvez, J. E. & Morte, A. A mycorrhizal helper bacterium alleviates drought stress in mycorrhizal Helianthemum almeriense plants by regulating water relations and plant hormones. Environ. Exp. Bot. 207, 105228 (2023).

    Google Scholar 

19. Berrios, L. et al. Positive interactions between mycorrhizal fungi and bacteria are widespread and benefit plant growth. Curr. Biol. 33, 2878–2887.e2874 (2023).

    Google Scholar 

20. Navarro-Rodenas, A., Berna, L. M., Lozano-Carrillo, C., Andrino, A. & Morte, A. Beneficial native bacteria improve survival and mycorrhization of desert truffle mycorrhizal plants in nursery conditions. Mycorrhiza 26, 769–779 (2016).

    Google Scholar 

21. Jorgensen, K., Clemmensen, K. E., Wallander, H. & Lindahl, B. D. Do ectomycorrhizal exploration types reflect mycelial foraging strategies? New Phytol. 237, 576–584 (2023).

    Google Scholar 

22. Durán, P. et al. Microbial interkingdom interactions in roots promote Arabidopsis survival. Cell 175, 973–983.e914 (2018).

    Google Scholar 

23. Bertness, M. D. & Callaway, R. Positive interactions in communities. Trends Ecol. Evol. 9, 191–193 (1994).

    Google Scholar 

24. Hammarlund, S. P. & Harcombe, W. R. Refining the stress gradient hypothesis in a microbial community. Proc. Natl. Acad. Sci. USA 116, 15760–15762 (2019).

    Google Scholar 

25. Hernandez, D. J., David, A. S., Menges, E. S., Searcy, C. A. & Afkhami, M. E. Environmental stress destabilizes microbial networks. ISME J. 15, 1722–1734 (2021).

    Google Scholar 

26. Gao, C. et al. Co-occurrence networks reveal more complexity than community composition in resistance and resilience of microbial communities. Nat. Commun. 13, 3867 (2022).

    Google Scholar 

27. Berrios, L., Ansell, T. B., Dahlberg, P. D. & Peay, K. G. Standardizing experimental approaches to investigate interactions between bacteria and ectomycorrhizal fungi. FEMS Microbiol. Rev. 49, fuae035 (2025).

    Google Scholar 

28. Hao, Z. G. et al. Thresholds in aridity and soil carbon-to-nitrogen ratio govern the accumulation of soil microbial residues. Commun. Earth Environ. 2, 236 (2021).

    Google Scholar 

29. Miyamoto, Y., Maximov, T. C., Kononov, A. & Sugimoto, A. Soil propagule banks of ectomycorrhizal fungi associated with Larix cajanderi above the treeline in the Siberian Arctic. Mycoscience 63, 142–148 (2022).

    Google Scholar 

30. Shemesh, H., Bruns, T. D., Peay, K. G., Kennedy, P. G. & Nguyen, N. H. Changing balance between dormancy and mortality determines the trajectory of ectomycorrhizal fungal spore longevity over a 15-yr burial experiment. New Phytol. 238, 11–15 (2023).

    Google Scholar 

31. Williams, A., Sinanaj, B. & Hoysted, G. A. Plant-microbe interactions through a lens: tales from the mycorrhizosphere. Ann. Bot. 133, 399–412 (2024).

    Google Scholar 

32. Hoek, T. A. et al. Resource availability modulates the cooperative and competitive nature of a microbial cross-feeding mutualism. PLoS Biol. 14, e1002540 (2016).

    Google Scholar 

33. Piccardi, P., Vessman, B. & Mitri, S. Toxicity drives facilitation between 4 bacterial species. Proc. Natl. Acad. Sci. USA 116, 15979–15984 (2019).

    Google Scholar 

34. Lofgren, L. et al. Suillus: an emerging model for the study of ectomycorrhizal ecology and evolution. New Phytol. 242, 1448–1475 (2024).

    Google Scholar 

35. Voller, F., Ardanuy, A., Taylor, A. F. S. & Johnson, D. Maintenance of host specialisation gradients in ectomycorrhizal symbionts. New Phytol. 242, 1426–1435 (2024).

    Google Scholar 

36. Frey-Klett, P., Garbaye, J. & Tarkka, M. The mycorrhiza helper bacteria revisited. New Phytol. 176, 22–36 (2007).

    Google Scholar 

37. Tedersoo, L., Bahram, M. & Zobel, M. How mycorrhizal associations drive plant population and community biology. Science 367, eaba1223 (2020).

    Google Scholar 

38. Peng, L. et al. A facultative ectomycorrhizal association is triggered by organic nitrogen. Curr. Biol. 32, 5235–5249.e5237 (2022).

    Google Scholar 

39. Rasheed, Z. et al. Production of some secondary metabolites of antibiotic nature from mycorrhizal helper bacteria (MHB) associated with conifers. Pol. J. Environ. Stud. 33, 4325–4334 (2024).

    Google Scholar 

40. Singh, B. K. & Walker, A. Microbial degradation of organophosphorus compounds. FEMS Microbiol. Rev. 30, 428–471 (2006).

    Google Scholar 

41. Yang, F. et al. Genome-wide analysis reveals genetic potential for aromatic compounds biodegradation of Sphingopyxis. Biomed. Res. Int. 2020, 5849123 (2020).

    Google Scholar 

42. Zhao, Q. et al. Comparative genomic analysis of 26 Sphingomonas and Sphingobium strains: dissemination of bioremediation capabilities, biodegradation potential and horizontal gene transfer. Sci. Total Environ. 609, 1238–1247 (2017).

    Google Scholar 

43. Tong, S. et al. Characterization of a fungal competition factor: production of a conidial cell-wall-associated antifungal peptide. PLoS Pathog. 16, e1008518 (2020).

    Google Scholar 

44. Zhong, Z. X. et al. Differential gene expression profiling analysis in Pleurotus ostreatus during interspecific antagonistic interactions with Dichomitus squalens and Trametes versicolor. Fungal Biol. 121, 1025–1036 (2017).

    Google Scholar 

45. Mochizuki, K. et al. The ASCT/SCS cycle fuels mitochondrial ATP and acetate production in Trypanosoma brucei. Biochim. Biophys. Acta Bioenerg. 1861, 148283 (2020).

    Google Scholar 

46. Labbe, J. L., Weston, D. J., Dunkirk, N., Pelletier, D. A. & Tuskan, G. A. Newly identified helper bacteria stimulate ectomycorrhizal formation in Populus. Front. Plant Sci. 5, 579 (2014).

    Google Scholar 

47. Al Farraj, D. A., Alkufeidy, R. M., Alkubaisi, N. A. & Alshammari, M. K. Polynuclear aromatic anthracene biodegradation by psychrophilic Sphingomonas sp., cultivated with Tween-80. Chemosphere 263, 128115 (2021).

    Google Scholar 

48. Park, J. et al. Sulfur metabolism-mediated fungal glutathione biosynthesis is essential for oxidative stress resistance and pathogenicity in the plant pathogenic fungus Fusarium graminearum. mBio 15, e0240123 (2024).

49. Hanna, V. S., Abd El-Ghany, M. N., Ibrahim, M. I. M., Abdel-Rahman, T. M. & Tallima, H. Novel approaches to Mortierella alpina identification and arachidonic acid production optimization. ACS Omega 9, 34456–34463 (2024).

    Google Scholar 

50. Agerer, R. Exploration types of ectomycorrhizae: a proposal to classify ectomycorrhizal mycelial systems according to their patterns of differentiation and putative ecological importance. Mycorrhiza 11, 107–114 (2001).

    Google Scholar 

51. van der Linde, S. et al. Environment and host as large-scale controls of ectomycorrhizal fungi. Nature 558, 243–248 (2018).

    Google Scholar 

52. Põlme, S. et al. Biogeography of ectomycorrhizal fungi associated with alders (Alnus spp.) in relation to biotic and abiotic variables at the global scale. New Phytol. 198, 1239–1249 (2013).

    Google Scholar 

53. Reis, F. et al. Ectomycorrhizal fungal diversity and community structure associated with cork oak in different landscapes. Mycorrhiza 28, 357–368 (2018).

    Google Scholar 

54. Edwards, J. et al. Structure, variation, and assembly of the root-associated microbiomes of rice. Proc. Natl. Acad. Sci. USA 112, E911–E920 (2015).

    Google Scholar 

55. He, X. H., Yang, Y. Z. & Yuang, Z. L. Protocol for sampling of root and rhizosphere soils from trees in natural fields. Bio 101, e2003655 (2021).

    Google Scholar 

56. Beckers, B., Op De Beeck, M., Weyens, N., Boerjan, W. & Vangronsveld, J. Structural variability and niche differentiation in the rhizosphere and endosphere bacterial microbiome of field-grown poplar trees. Microbiome 5, 25 (2017).

    Google Scholar 

57. Chelius, M. K. & Triplett, E. W. The diversity of archaea and bacteria in association with the roots of Zea mays L. Microb. Ecol. 41, 252–263 (2001).

    Google Scholar 

58. Bodenhausen, N., Horton, M. W. & Bergelson, J. Bacterial communities associated with the leaves and the roots of Arabidopsis thaliana. PloS ONE 8, e56329 (2013).

    Google Scholar 

59. Gardes, M. & Bruns, T. D. ITS primers with enhanced specificity for basidiomycetes-application to the identification of mycorrhizae and rusts. Mol. Ecol. 2, 113–118 (1993).

    Google Scholar 

60. White, T. J., Bruns, T., Lee, S. & Taylor, J. Amplification and direct sequencing of fungal ribosomal RNA genes for phylogenetics. PCR Protocols. https://doi.org/10.1016/b978-0-12-372180-8.50042-1 (Academic Press, 1990).

61. Põlme, S. et al. FungalTraits: a user-friendly traits database of fungi and fungus-like stramenopiles. Fungal Divers. 105, 1–16 (2021).

    Google Scholar 

62. Kim, M., Oh, H. S., Park, S. C. & Chun, J. Towards a taxonomic coherence between average nucleotide identity and 16S rRNA gene sequence similarity for species demarcation of prokaryotes. Int. J. Syst. Evol. Microbiol. 64, 346–351 (2014).

    Google Scholar 

63. Deveau, A. et al. The mycorrhiza helper Pseudomonas fluorescens BBc6R8 has a specific priming effect on the growth, morphology and gene expression of the ectomycorrhizal fungus Laccaria bicolor S238N. New Phytol. 175, 743–755 (2007).

    Google Scholar 

64. Zhao, L., Wu, X. Q., Ye, J. R., Li, H. & Li, G. E. Isolation and characterization of a mycorrhiza helper bacterium from rhizosphere soils of poplar stands. Biol. Fertil. Soils 50, 593–601 (2014).

    Google Scholar 

65. Founoune, H. et al. Interactions between ectomycorrhizal symbiosis and Fluorescent pseudomonads on Acacia holosericea: isolation of mycorrhiza helper bacteria (MHB) from a Soudano-Sahelian soil. FEMS Microbiol. Ecol. 41, 37–46 (2002).

    Google Scholar 

66. Zi, H. Y. et al. Co-inoculation with rhizobacterial community and an ectomycorrhizal fungus promotes poplar ectomycorrhization. Appl. Soil Ecol. 206, 105833 (2025).

    Google Scholar 

67. Yu, L. et al. Rhizosphere microbiome of forest trees is connected to their resistance to soil-borne pathogens. Plant Soil 479, 143–158 (2022).

    Google Scholar 

68. Cavanaugh, J. E. & Neath, A. A. The Akaike information criterion: background, derivation, properties, application, interpretation, and refinements. Wiley Interdiscip. Rev. Comput. Stat. 11, e1460 (2019).

69. Muggeo, V. Segmented: an R package to fit regression models with broken-line relationships. R New. 8, 20–25 (2008).

70. Hu, W. G. et al. Aridity-driven shift in biodiversity-soil multifunctionality relationships. Nat. Commun. 12, 5350 (2021).

    Google Scholar 

71. Friedman, J. & Alm, E. J. Inferring correlation networks from genomic survey data. PLoS Comput. Biol. 8, e1002687 (2012).

    Google Scholar 

72. Herren, C. M. & McMahon, K. D. Cohesion: a method for quantifying the connectivity of microbial communities. ISME J. 11, 2426–2438 (2017).

    Google Scholar 

73. Dufrêne, M. & Legendre, P. Species assemblages and indicator species: the need for a flexible asymmetrical approach. Ecol. Monogr. 67, 345–366 (1997).

    Google Scholar 

74. Gweon, H. S. et al. Contrasting community assembly processes structure lotic bacteria metacommunities along the river continuum. Environ. Microbiol. 23, 484–498 (2021).

    Google Scholar 

Download references