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Aims Decomposition of leaf litter is influenced by litter quality as determined by plant genotype and environment, as well as climate and soil properties. We studied these drivers of decomposition in communities of Salix varieties, hypothesizing that decomposition rates would increase under warmer climate, in more diverse communities, and with increasing litter quality of the individual varieties. Methods Litter from four Salix varieties was incubated in three field trials across a latitudinal gradient from Central to Northern Europe. Litter and stand properties were measured and used as predictors of decomposition. Results No significant site differences in remaining mass or nitrogen were found. Instead, effects of initial leaf litter quality on decomposition were stronger than climatic effects. Litter quality of individual varieties strongly affected decomposition, while increasing litter diversity did not. Conclusions Decomposition was controlled by variety identity depending on site, indicating that local soil conditions affect litter quality (and thus decomposition) more than macroclimate. In mixed communities, varieties producing fast-decomposing litter enhanced the litter decomposition of other components producing slow-decomposing litter, and vice versa. This implies that site conditions partly determine which varieties affect community-level decomposition and nutrient release.

While it is well established that plants associating with arbuscular mycorrhizal (AM) and ectomycorrhizal (ECM) fungi cycle carbon (C) and nutrients in distinct ways, we have a limited understanding of whether varying abundance of ECM and AM plants in a stand can provide integrative proxies for key biogeochemical processes. We explored linkages between the relative abundance of AM and ECM trees and microbial functioning in three hardwood forests in southern Indiana, USA. Across each site’s ‘mycorrhizal gradient’, we measured fungal biomass, fungal: bacterial (F: B) ratios, extracellular enzyme activities, soil carbon: nitrogen ratio, and soil pH over a growing season. We show that the percentage of AM or ECM trees in a plot promotes microbial communities that both reflect and determine the C to nutrient balance in soil. Soils dominated by ECM trees had higher F: B ratios and more standing fungal biomass than AM stands. Enzyme stoichiometry in ECM soils shifted to higher investment in extracellular enzymes needed for nitrogen and phosphorus acquisition than in C-acquisition enzymes, relative to AM soils. Our results suggest that knowledge of mycorrhizal dominance at the stand or landscape scale may provide a unifying framework for linking plant and microbial community dynamics, and predicting their effects on ecological function.

Although much is known about how trees and their associated microbes influence nitrogen cycling in temperate forest soils, less is known about biotic controls over phosphorus (P) cycling. Given that mycorrhizal fungi are instrumental for P acquisition and that the two dominant associations – arbuscular mycorrhizal (AM) and ectomycorrhizal (ECM) fungi – possess different strategies for acquiring P, we hypothesized that P cycling would differ in stands dominated by trees associated with AM vs ECM fungi. We quantified soil solution P, microbial biomass P, and sequentially extracted inorganic and organic P pools from May to November in plots dominated by trees forming either AM or ECM associations in south‐central Indiana, USA. Overall, fungal communities in AM and ECM plots were functionally different and soils exhibited fundamental differences in P cycling. Organic forms of P were more available in ECM plots than in AM plots. Yet inorganic P decreased and organic P accumulated over the growing season in both ECM and AM plots, resulting in increasingly P‐limited microbial biomass. Collectively, our results suggest that P cycling in hardwood forests is strongly influenced by biotic processes in soil and that these are driven by plant‐associated fungal communities.

Better understanding and quantifying the relative influence of plants, associated mycorrhizal fungi, and abiotic factors such as elevatedCO2 on biotic weathering is essential to constraining weathering estimates. We employed a column microcosm system to examine the effects of elevated CO2 and Pinus sylvestris seedlings, with or without the ectomycorrhizal fungiPiloderma fallax and Suillus variegatus, on rhizosphere soil solution concentrations of low-molecular-weight organic acids (LMWOAs) and on the weathering of primary minerals. Seedlings significantly increased mineral weathering, as estimated from elemental budgets of Ca, K, Mg, and Si. Elevated CO2 increased plant growth and LMWOA concentrations but had no effect on weathering. Colonization by ectomycorrhizal fungi, particularly P. fallax, showed some tendency to increase weathering. LMWOA concentrations correlated with seedling biomass across bothCO2 and mycorrhizal treatments but not with total weathering. We conclude that nutrient uptake, which reduces transport limitation to weathering, is the primary mechanism by which plants enhanced weathering in this system. While the experimental system used departs from conditions in forest soils in a number of ways, these results are in line with weathering studies performed at the ecosystem, macrocosm, and microcosm scale, indicating that nutrient uptake by plants and microbes is an important biological mechanism by which mineral weathering is enhanced.

Willows (Salix spp.) are mycorrhizal tree species sometimes cultivated as short rotation coppice (SRC) on arable sites for energy purposes; they are also among the earliest plants colonising primary successional sites in natural stands. The objective of this study was to analyse the degree of colonisation and diversity of ectomycorrhizal (EM) communities on willows grown as SRC in arable soils and their adjacent natural or naturalized stands. Arable sites usually lack ectomycorrhizal host plants before the establishment of SRC, and adjacent natural or naturalized willow stands were hypothesized to be a leading source of ectomycorrhizal inoculum for the SRC. Three test sites including SRC stands (Salix viminalis, Salix dasyclados, and Salix schwerinii) and adjacent natural or naturalized (Salix caprea, Salix fragilis, and Salix × mollissima) stands in central Sweden were investigated on EM colonisation and morphotypes, and the fungal partners of 36 of the total 49 EM fungi morphotypes were identified using molecular tools. The frequency of mycorrhizas in the natural/naturalized stands was higher (two sites) or lower (one site) than in the corresponding cultivated stands. Correspondence analysis revealed that some EM taxa (e.g. Agaricales) were mostly associated with cultivated willows, while others (e.g. Thelephorales) were mostly found in natural/naturalized stands. In conclusion, we found strong effects of sites and willow genotype on EM fungi formation, but poor correspondence between the EM fungi abundance and diversity in SRC and their adjacent natural/naturalized stands. The underlying mechanism might be selective promotion of some EM fungi species by more effective spore dispersal.

Root and mycelial exudation contributes significantly to soil carbon (C) fluxes, and is likely to be altered by an elevated atmospheric carbon dioxide (CO₂) concentration and nitrogen (N) deposition. We quantified soluble, low-molecular-weight (LMW) organic compounds exuded by ectomycorrhizal plants grown under ambient (360 p.p.m.) or elevated (710 p.p.m.) CO₂ concentrations and with different N sources. Scots pine seedlings, colonized by one of five different ectomycorrhizal or nonmycorrhizal fungi, received 70 μM N, either as NH₄Cl or as alanine, in a liquid growth medium. Exudation of LMW organic acids (LMWOAs), dissolved monosaccharides and total dissolved organic carbon were determined. Both N and CO₂ had a significant impact on exudation, especially of LMWOAs. Exudation of LMWOAs was negatively affected by inorganic N and decreased by 30-85% compared with the organic N treatment, irrespective of the CO₂ treatment. Elevated CO₂ had a clear impact on the production of individual LMWOAs, although with very contrasting effects depending on which N source was supplied.

We examined exudation of low molecular weight (LMW) organic compounds of ectomycorrhizal (ECM) and non-mycorrhizal (NM) seedlings in relation to metals. Scots pine seedlings, either colonized by one of six different ECM fungi or NM, were grown in Petri dishes containing glass beads and liquid growth medium and exposed to elevated concentrations of Pb, Cd and As. Exudation of LMW organic compounds (LMW organic acids (LMWOAs), amino acids and dissolved monosaccharides) and dissolved organic carbon (DOC) was determined qualitatively and quantitatively and exudation rates were calculated. Metals had a significant impact on exudation, especially of oxalate. For Pb and Cd treatments, exudation of oxalate and total LMWOAs generally increased by 15–45% compared to nutrient controls. Production of amino acids, dissolved monosaccharides and DOC was not significantly stimulated by exposure to metals; however, there were non-significant trends towards increased exudation. Finally, exudation generally increased in the presence of mycorrhizal seedlings compared to NM seedlings. The results suggest that ECM fungi may reduce the toxicity of metals to plants through significant increases in the production of organic chelators. Axenic conditions are required to assess the full potential for production of these molecules but their overall significance in soil ecosystems needs to be determined using additional experiments under more ecologically realistic conditions.

The elimination of hazardous compounds in chemical wastes can be a complex and technically demanding task. In the search for environmental-friendly technologies, fungal mediated remediation and removal procedures are of concern. In this study, we investigated whether there are fungal species that can survive and grow on solely amine-containing compounds. One compound containing a primary amine group; 2-diethylaminoethanol, one compound with a primary amide group; 2,6-dichlorobenzamide (BAM), and a third compound containing a quaternary ammonium group; N 3 -trimethyl(2-oxiranyl)methanaminium chloride, were selected. The choice of these compounds was motivated by their excessive use in large scale manufacturing of protein separation media (2-diethylaminoethanol and the quaternary amine). 2,6-dichlorobenzamide, the degradation product of the herbicide 2,6-dichlorobenzonitrile (dichlobenil), was chosen since it is an extremely recalcitrant compound. Utilising part of the large fungal diversity in Northern European forests, a screening study using 48 fungal isolates from 42 fungal species, including saprotrophic and mycorrhizal fungi, was performed to test for growth responses to the chosen compounds. The ericoid (ERM) mycorrhizal fungus Rhizoscyphus ericae showed the best overall growth on 2-diethylaminoethanol and BAM in the 1–20 g L -1 concentration range, with a 35-fold and 4.5-fold increase in biomass, respectively. For N 3 -trimethyl(2-oxiranyl)methanaminium chloride, the peak growth occurred at 1 g L -1 . In a second experiment, including three of the most promising fungi ( Laccaria laccata , Hygrophorus camarophyllus and Rhizoscyphus ericae ) from the screening experiment, a simulated process water containing 1.9% (w/v) 2-diethylaminoethanol and 0.8% (w/v) N 3- trimethyl(2-oxiranyl)methanaminium chloride was used. Laccaria laccata showed the best biomass increase (380%) relative to a control, while the accumulation for Rhizoscyphus ericae and Hygrophorus camarophyllus were 292% and 136% respectively, indicating that mycorrhizal fungi can use amine- and amide-containing substrates as nutrients. These results show the potential of certain fungal species to be used in alternative green wastewater treatment procedures.

In boreal forest soils, ectomycorrhizal fungi are fundamentally important for carbon (C) dynamics and nutrient cycling. Although their extraradical mycelium (ERM) is pivotal for processes such as soil organic matter build-up and nitrogen cycling, very little is known about its dynamics and regulation. In this study, we quantified ERM production and turnover, and examined how these two processes together regulated standing ERM biomass in seven sites forming a chronosequence of 12- to 100-yr-old managed Pinus sylvestris forests. This was done by determining ERM biomass, using ergosterol as a proxy, in sequentially harvested in-growth mesh bags and by applying mathematical models. Although ERM production declined with increasing forest age from 1.2 to 0.5 kg ha−1 d−1, the standing biomass increased from 50 to 112 kg ha−1. This was explained by a drastic decline in mycelial turnover from seven times to one time per year with increasing forest age, corresponding to mean residence times from 25 d up to 1 yr. Our results demonstrate that ERM turnover is the main factor regulating biomass across differently aged forest stands. Explicit inclusion of ERM parameters in forest ecosystem C models may significantly improve their capacity to predict responses of mycorrhiza-mediated processes to management and environmental changes.

Societal Impact Statement Efficient mitigation of climate change requires predictive models of forest ecosystems as sinks for atmospheric carbon. Mycorrhizal fungi are drivers of soil carbon storage in boreal forests, yet they are typically excluded from ecosystem models, because of a lack of information about their growth and turnover. Closing this knowledge gap could help us better predict future responses to climate change and guide policy decisions for sustainable management of forest ecosystems. This study provides new estimates of the production and turnover of mycorrhizal mycelial biomass and necromass. This information can facilitate the integration of mycorrhizal fungi into new predictive models of boreal forest soils. Summary In boreal forests, turnover of biomass and necromass of ectomycorrhizal extraradical mycelia (ERM) are important for mediating long‐term carbon storage. However, ectomycorrhizal fungi are usually not considered in ecosystem models, because data for parameterization of ERM dynamics is lacking. Here, we estimated the production and turnover of ERM biomass and necromass across a hemiboreal Pinus sylvestris chronosequence aged 12 to 100 years. Biomass and necromass were quantified in sequentially harvested in‐growth bags, and incubated in the soil for 1–24 month, and Bayesian calibration of mathematical models was applied to arrive at parametric estimates of ERM production and turnover rates of biomass and necromass. Steady states were predicted to be nearly reached after 160 and 390 growing season days, respectively, for biomass and necromass. The related turnover rates varied with 95% credible intervals of 1.7–6.5 and 0.3–2.5 times yr−1, with mode values of 2.9 and 0.9 times yr−1, corresponding to mean residence times of 62 and 205 growing season days. Our results highlight that turnover of necromass is one‐third of biomass. This together with the variability in the estimates can be used to parameterize ecosystem models, to explicitly include ERM dynamics and its impact on mycorrhizal‐derived soil carbon accumulation in boreal forests. Efficient mitigation of climate change requires predictive models of forest ecosystems as sinks for atmospheric carbon. Mycorrhizal fungi are drivers of soil carbon storage in boreal forests, yet they are typically excluded from ecosystem models, because of a lack of information about their growth and turnover. Closing this knowledge gap could help us better predict future responses to climate change and guide policy decisions for sustainable management of forest ecosystems. This study provides new estimates of the production and turnover of mycorrhizal mycelial biomass and necromass. This information can facilitate the integration of mycorrhizal fungi into new predictive models of boreal forest soils.