The mycorrhizal tragedy of the commons
Abstract
- trees trade C for N with mycorrhizal network
- this network is the “commons”, part of the tragedy
- trees gain additional N et the expense of neighbours by supplying more C to fungi
- this can lead to increased N immobilization (in fungal biomass)
Introduction
- ectomycorrhizal fungi (EMF) hold N back from trees when they are in need
- when N supply is amended by fertilization, also EMF give away greater proportion
- further supply of EMF with C makes them diminish N returns
- fungus competes with other EMF symbionts of the same plant, can gain more C by exporting more N, until personal N matches its received C
- receiving more C makes EMF use more N for itself
- enhanced EMF growth initially increases N uptake, but eventually N immobilization in fungal biomass; negative feedback on plant’s N uptake
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How can this negative feedback made it through evolution?
- dual scale of ectomycorrhizal symbiosis: one tree, many fungi and many trees, one fungus
- trees have optimize individual benefit, this has consequences for other trees connected through the fungal network
- combined efforts of trees to increase individual N gain aggravates N immobilization in fungal biomass
Hypothesis
- fungus shares greater N proportion with trees that deliver more C
- tree delivers more C to fungi that supply more N
- tragedy from tree viewpoint: total C export to all fungi so high that it leads to N immobilization
- tragedy from fungus viewpoint: exporting more N would reduce own growth, exporting less N reducec competitiveness for plant C
- plants share common N pool, optimizing for individual gain hampers comunity gain
- experiments with plant strangling and shading
Materials and methods
Seedling experiment
Field experiment
Model description
Results
- shaded less biomass than sun, strangled no effect
- shaded plants more N than sun plants, strangled plants way less: decreased C export to fungi mobilizes soil N to the plant and vice versa
- maximum network-scale N mobilization occurs at intermediate level of network-scale C export
- at individual tree level: 1 more C supply leads to 0.95 more N return, hence there is competition among trees which will eventually make network-scale N mobilization drop
Discussion
- tragedy of the commons: individual gain reduces community gain
- there is one common N pool all trees draw from (common EMF network, in which multiple fungi connect the host plants)
- elevated atmospheric CO$_2$ levels should lead to higher N immobilization, more fungal biomass growth, maybe less tree growth