Science · M3

Soil Biodiversity

Beneath every thriving guild lies two invisible communities working in parallel — fungi expanding every root's reach into the soil, and bacteria fixing the very air into food for your plants.

Fungi

Mycorrhizal partnerships and fungal disease control

Bacteria

Nitrogen-fixing Rhizobium and free-living fixers

This partnership is older than trees. The earliest fossils of mycorrhizal roots date to 407 million years ago, placing them very early in land-plant history (Brundrett & Tedersoo 2018). Over 80% of land plants — more than 250,000 species — still form these associations today (Martin & van der Heijden 2024). They come in four distinct forms: arbuscular (AM), ectomycorrhizal (EcM), ericoid, and orchid mycorrhizas, each built from different fungi with different root architectures. Only about 10% of plants go it entirely alone.

What the fungi provide is remarkable in its quiet scale. Up to 90% of a plant's phosphorus and nitrogen can arrive through mycorrhizal hyphae rather than through the roots themselves (van der Heijden et al. 2008). In return, plants channel 5–30% of their photosynthate underground to sustain the fungal network — a flow so large that globally, the carbon entering mycorrhizal mycelium equals roughly 36% of annual fossil fuel CO₂ emissions (Hawkins et al. 2023). The hyphae also bind soil particles into aggregates, building the physical structure of the earth beneath your feet.

Soil cross-section showing two small green plants connected beneath the ground by an extensive network of fine white fungal hyphae (mycorrhizal mycelium) threading through dark earth
Fine white fungal hyphae threading through the soil around two seedlings, extending far beyond what the roots alone could reach. In many plant communities, this is not a rare special case but a common part of how plants feed.

The Mycorrhizal Balance

Luo et al. (2023) analysed 74,563 forest inventory plots across 35 US ecoregions and found that in ~77% of ecoregions, forests mixing arbuscular mycorrhizal (AM) and ectomycorrhizal (EcM) host species were more productive than forests dominated by either type alone. The mixing benefit was most pronounced at low species richness (≤5 species) — the range most directly comparable to designed plant guilds.

Side-by-side botanical illustration comparing ectomycorrhizal (EcM) and arbuscular mycorrhizal (AM) root anatomy. Left: EcM — a blue-violet fungal mantle sheaths the root exterior, with the Hartig net growing between cells only, never entering them. Right: AM — a golden trunk hypha threads between cells, sending branches inside each cell where they form tree-like arbuscules, with vesicles and an external spore visible.
Left — EcM fungi wrap the root in a dense mantle, with the Hartig net growing between cells only — never entering them. This extracellular network unlocks organic nitrogen from leaf litter. Right — AM fungi thread a trunk hypha between cells, sending branches inside each cell where they form tree-like arbuscules — the exchange surfaces for phosphorus from mineral soil. Their core functions are largely non-overlapping.

In tropical gardens: most plants are AM hosts by default. To activate the mixing benefit you must deliberately include EcM-associating species — dipterocarps (meranti, shorea), tropical oaks, or Casuarina. Without at least one, the mycorrhizal mixing component contributes nothing to your score.

Mycorrhizal type assignments follow Brundrett & Tedersoo (2018), which compiled the family-level mycorrhizal classification used globally.

TypeCommon familiesSoil layerNutrient form
AMMost herbs, tropical trees, cropsMineralSoluble P, mineralised N
EcMPinaceae, Fagaceae, Betulaceae, DipterocarpaceaeOrganicOrganic N & P (enzymatic)
NMBrassicaceae, Chenopodiaceae, ProteaceaeNo mycorrhizal partner

Mycoparasitic Disease Suppression

Trichoderma, Clonostachys, and related fungal species are the soil's own disease fighters — free-living fungi that hunt and dissolve soil-borne pathogens. Unlike mycorrhizal fungi, they don't form nutrient-exchange partnerships — but they can colonize root surfaces, trigger plant defences, and reduce the population of disease-causing fungi around your plants' roots. When Trichoderma strains are deliberately applied as inoculants, pathogen reductions of 5–87% have been documented depending on the strain, target pathogen, and host crop — with the strongest results in this review against Fusarium species and root-rot pathogens (Singh et al. 2024, a review of biocontrol studies).

Critically, Trichoderma discriminates between beneficial fungi and pathogens at a distance (Stange et al. 2024) — growing toward pathogens while avoiding ectomycorrhizal fungi in laboratory assays. A guild designed for rich mycorrhizal connections and one with active Trichoderma aren't in conflict — they coexist without interference.

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Chemical detection at a distance

Trichoderma detects fungal pathogens at a distance — before any physical contact. It grows toward pathogens while showing the opposite response to ectomycorrhizal fungi. This targeted response is why it behaves more like a biocontrol agent than a blanket antifungal (Stange et al. 2024).

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Attack and dissolve

Trichoderma secretes chitinases and glucanases — enzymes that break down the cell walls of pathogens like Fusarium, Rhizoctonia, and Pythium. Singh et al. (2024) documented up to 87% pathogen reduction, though efficacy varies widely by strain and target.

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Spares mycorrhizal networks

In laboratory assays, Trichoderma actively grew away from EcM fungi and did not activate its mycoparasitic genes against them — reserving its dissolving enzymes for pathogens (Stange et al. 2024). This suggests disease suppression and mycorrhizal diversity can coexist, though field confirmation is still limited.

Nitrogen-Fixing Bacteria

Nitrogen is often a limiting nutrient in plant growth. Plants cannot use atmospheric nitrogen directly. Nitrogen-fixing bacteria solve this problem by breaking the N≡N triple bond and converting atmospheric nitrogen into ammonium (NH₄⁺) that plants can absorb.

The most important partnership in a garden guild is between legumes (Fabaceae) and Rhizobium bacteria. The plant provides carbon-rich photosynthate to fuel the bacterium; the bacterium provides fixed nitrogen in return. Root nodules are the physical site of this exchange. Inside them, rhizobial bacteroids use nitrogenase to convert atmospheric N₂ into ammonia, which the plant then turns into ammonium (NH₄⁺). Leghemoglobin helps keep oxygen levels low enough for that reaction to work.

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Legume–Rhizobium

The dominant pathway. Many Fabaceae (beans, peas, clover, acacia, soy) form nodules with specific Rhizobium strains, and the exact outcome depends on the host, the bacterial strain, and the soil environment.

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Non-Legume Symbiosis

Frankia bacteria partner with alder (Alnus) and casuarinas. Less common than legume nodules, but still important in some woody lineages. Allocasuarina is unusual — it also forms EcM associations (Brundrett & Tedersoo 2018), giving it dual soil benefits.

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Free-Living Fixers

This page's score focuses most strongly on the better-studied symbioses with legumes and actinorhizal hosts. Free-living nitrogen fixers are not the main driver of this metric.

Why legumes score so high: legumes can contribute nitrogen to the guild over time — shared through root turnover, nodule decomposition, and movement into the surrounding soil.

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Infection and nodule formation

Rhizobium bacteria in the soil detect flavonoid signals released by legume roots. The bacteria enter through the root hairs, forming an infection thread that travels deep into the root where nodules develop.

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Nitrogen fixation

Inside nodules, bacteroids express nitrogenase — the enzyme that breaks the N≡N bond. Leghemoglobin helps regulate oxygen levels so the anaerobic reaction can proceed. The product is NH₄⁺.

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Nitrogen release to the guild

Fixed nitrogen enters the soil as roots leak it, as old nodules break down, and as fallen leaves decay. Neighbouring plants absorb this nitrogen directly — free fertiliser for every plant sharing the same soil volume.

How legumes share nitrogen with neighbours through mycorrhizal networks: nitrogen fixation at nodules and transfer through a common mycorrhizal network (CMN)
The legume–fungi synergy. Rhizobium bacteria fix nitrogen inside root nodules (1). Mycorrhizal hyphae then form a common network (CMN) connecting the legume's roots to its neighbours (2). Our synthesis of ¹⁵N isotope-tracing data found that nitrogen transfer to neighbours was 12.6× higher when both mycorrhizal network and N-fixer were present versus neither alone — and 31% above what either mechanism would predict additively. However, whether this nitrogen translates into measurable biomass gains depends heavily on context: a separate meta-analysis of 31 CMN studies found mostly neutral effects on plant biomass (Lehmann et al. 2025), suggesting the pipe works but downstream responses vary with context.

How We Score It

Your soil biodiversity score combines three things: whether your guild hosts both types of mycorrhizal fungi, how many nitrogen-fixers it includes, and whether any plants are known hosts of disease-fighting fungi.

40%
AM:EcM Balance
Do your plants host both types of soil fungi? A guild with only AM hosts (or only EcM hosts) scores zero for this component. The best score goes to a roughly even mix — inspired by Luo et al. (2023), who found that forests mixing both types were more productive than forests dominated by either one alone.
Score Pure AM 50:50 mix Pure EcM best score

The scoring curve peaks at a 50:50 AM:EcM mix. The exact optimum isn't specified in the paper — we use 50:50 because both fungal types access different nutrients in different soil layers, and their benefits are largely complementary.

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40%
N-Fixation Capacity
What share of your guild can fix nitrogen from the air? More fixers means more free nitrogen entering the soil. Adding even one legume helps — and the benefit keeps growing as you add more, though with diminishing returns. Handa et al. (2014) showed that combining N-fixer litter with fast-decomposing litter increased nitrogen release by 32.5%.
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20%
Mycoparasite Capacity
Do any of your plants host disease-fighting fungi like Trichoderma? When applied as inoculants, these fungi have reduced soil pathogens by 5–87% depending on strain and target (Singh et al. 2024). More known hosts in your guild means a higher score.

What We Can't Know Yet

Temporal lag. Fungal communities assemble over years. Bacterial nodule formation requires the right Rhizobium strain to be present in your soil. Scores represent medium-term potential, not day-one function.
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Rhizobium strain specificity. Different legume species require specific Rhizobium strains. If the matching strain is absent from your soil, nodulation may be weak or fail.
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Mycoparasite data is sparse. Our query of the GloBI database finds mycoparasite associations for ~0.4% of plant species. We reward known hosts only, so this part of the score is limited by sparse records.
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Legumes are a pathogen exception. Wang et al. (2023) found positive plant-soil feedback for Fabaceae (95% CI 0.23–0.89), consistent with limited pathogen specialization within the family and specialist nitrogen-fixing rhizobia. In practice, that means the usual pathogen-dilution story may work differently for legumes.

References

Luo, Phillips, Jo et al. 2023 Higher productivity in forests with mixed mycorrhizal strategies. Nat. Commun. 14, 7654. doi:10.1038/s41467-023-36888-0 Wang, Burrill, Podzikowski et al. 2023 Dilution of specialist pathogens drives productivity benefits from diversity in plant mixtures. Nat. Commun. 14, 2881. doi:10.1038/s41467-023-44253-4 Xu, Zheng, Liu et al. 2025 Fungal biodiversity buffers the impacts of plant diversity loss on soil ecosystem multifunctionality. Nat. Commun. 16, 1847. doi:10.1038/s41467-025-60661-0 Handa, Aerts, Berendse et al. 2014 Consequences of biodiversity loss for litter decomposition across biomes. Nature 509, 218–221. doi:10.1038/nature13247 Wagg, Bender, Widmer & van der Heijden 2014 Soil biodiversity and soil community composition determine ecosystem multifunctionality. PNAS 111, 5266–5270. doi:10.1073/pnas.1320054111 Brundrett & Tedersoo 2018 Evolutionary history of mycorrhizal symbioses and global host plant diversity. New Phytol. 220, 1108–1115. doi:10.1111/nph.14976 Stange, Kersting, Padmanaban et al. 2024 The decision for or against mycoparasitic attack by Trichoderma spp. is taken already at a distance in a prey-specific manner. Fungal Biol. Biotechnol. doi:10.1186/s40694-024-00183-4 Singh, Singh, Pradhan et al. 2024 Harnessing Trichoderma mycoparasitism as a tool in the management of soil dwelling plant pathogens. Microb. Ecol. 87, 158. doi:10.1007/s00248-024-02472-2 Martin & van der Heijden 2024 The mycorrhizal symbiosis: research frontiers in genomics, ecology, and agricultural application. New Phytol. 242, 1486–1506. doi:10.1111/nph.19541 Hawkins, Cargill, Aponte et al. 2023 Mycorrhizal mycelium as a global carbon pool. Curr. Biol. 33, R560–R573. doi:10.1016/j.cub.2023.02.027 van der Heijden, Bardgett & van Straalen 2008 The unseen majority: soil microbes as drivers of plant diversity and productivity in terrestrial ecosystems. Ecol. Lett. 11, 296–310. doi:10.1111/j.1469-8137.2008.02681.x Herridge, Peoples & Boddey 2008 Global inputs of biological nitrogen fixation in agricultural systems. Plant Soil 311, 1–18. doi:10.1007/s11104-008-9668-3 Lehmann, Tang & Rillig 2025 Meta-analysis of effects of common mycorrhizal networks formed by arbuscular mycorrhizal fungi on plant and fungal parameters. Funct. Ecol. doi:10.1111/1365-2435.70125