• Energy Research

Large-scale liming and wood ash addition are common practices to mitigate soil and water acidification in temperate and boreal forests. In addition, wood ash recycles nutrients removed at harvest to the forest ecosystem. Both liming and wood ash applications typically increase soil pH by 1–2 units. Therefore, they affect a range of soil processes and organisms including ectomycorrhizal (EM) fungi which are vital for the nutrition of many tree species. Here we review field studies reporting the effects of lime and wood ash amendments on EM fungi. We systematically compiled studies where known amounts of ash or lime were distributed to plots paired with comparable control plots, and where mycorrhizal variables were recorded. For a subset of studies meeting explicit criteria, we performed meta-analyses using overall mycorrhizal abundance, species richness or the abundance of specific fungal taxa as response variables. Guided by availability of data, the focus is on Nordic coniferous forests. Although the reviewed field studies varied widely in dosage and experimental setup they clearly demonstrated that liming and wood ash amendments influence EM fungal species composition. Across studies, species belonging to the lineages Cortinarius and Russula-lactarius (Basidiomycota), particularly Russula ochroleuca, decreased in relative abundance, while species within the Tuber-helvella (Ascomycota) increased. Particular species within the Amphinema-tylospora lineage responded in opposite directions; Tylospora fibrillosa decreased in relation to the control, while Amphinema byssoides increased. The significant changes in species or clade abundances were in the range of 5–20% compared to non-treated plots. In contrast, neither the belowground mycorrhizal biomass nor species richness responded to liming or wood ash applications. We conclude that liming and wood ash amendments cause consistent EM fungal species dominance shifts, but that a high EM fungal biomass and species and phylogenetic richness is maintained on the tree roots. Given the large dispersal potential of many EM fungi, we therefore suggest that these treatments at normal recommended dosages do not pose any immediate threats to EM fungal biodiversity, at least not when applied at relatively small spatial scales. Whether the observed dominance shifts among EM fungal clades have consequences for the functioning of the EM fungal guild, e.g. in relation to nutrient cycling or tree nutrition, is an important question that should be further investigated.

Summary Ectomycorrhizal fungi are essential for nitrogen (N) cycling in many temperate forests and responsive to anthropogenic N addition, which generally decreases host carbon (C) allocation to the fungi. In the boreal region, however, ectomycorrhizal fungal biomass has been found to correlate positively with soil N availability. Still, responses to anthropogenic N input, for instance through atmospheric deposition, are commonly negative. To elucidate whether variation in N supply affects ectomycorrhizal fungi differently depending on geographical context, we investigated ectomycorrhizal fungal communities along fertility gradients located in two nemo‐boreal forest regions with similar ranges in soil N : C ratios and inorganic N availability but contrasting rates of N deposition. Ectomycorrhizal biomass and community composition remained relatively stable across the N gradient with low atmospheric N deposition, but biomass decreased and the community changed more drastically with increasing N availability in the gradient subjected to higher rates of N deposition. Moreover, potential activities of enzymes involved in ectomycorrhizal mobilisation of organic N decreased as N availability increased. In forests with low external input, we propose that stabilising feedbacks in tree‐fungal interactions maintain ectomycorrhizal fungal biomass and communities even in N‐rich soils. By contrast, anthropogenic N input seems to impair ectomycorrhizal functions.

AbstractArctic and alpine tundra ecosystems are large reservoirs of organic carbon1,2. Climate warming may stimulate ecosystem respiration and release carbon into the atmosphere3,4. The magnitude and persistency of this stimulation and the environmental mechanisms that drive its variation remain uncertain5–7. This hampers the accuracy of global land carbon–climate feedback projections7,8. Here we synthesize 136 datasets from 56 open-top chamber in situ warming experiments located at 28 arctic and alpine tundra sites which have been running for less than 1 year up to 25 years. We show that a mean rise of 1.4 °C [confidence interval (CI) 0.9–2.0 °C] in air and 0.4 °C [CI 0.2–0.7 °C] in soil temperature results in an increase in growing season ecosystem respiration by 30% [CI 22–38%] (n = 136). Our findings indicate that the stimulation of ecosystem respiration was due to increases in both plant-related and microbial respiration (n = 9) and continued for at least 25 years (n = 136). The magnitude of the warming effects on respiration was driven by variation in warming-induced changes in local soil conditions, that is, changes in total nitrogen concentration and pH and by context-dependent spatial variation in these conditions, in particular total nitrogen concentration and the carbon:nitrogen ratio. Tundra sites with stronger nitrogen limitations and sites in which warming had stimulated plant and microbial nutrient turnover seemed particularly sensitive in their respiration response to warming. The results highlight the importance of local soil conditions and warming-induced changes therein for future climatic impacts on respiration.

Abstract Changes in fire regime of boreal forests in response to climate warming are expected to impact postfire recovery. However, quantitative data on how managed forests sustain and recover from recent fire disturbance are limited. Two years after a large wildfire in managed even‐aged boreal forests in Sweden, we investigated how recovery of aboveground and belowground communities, that is, understory vegetation and soil microbial and faunal communities, responded to variation in the severity of soil (i.e., consumption of soil organic matter) and canopy fires (i.e., tree mortality). While fire overall enhanced diversity of understory vegetation through colonization of fire adapted plant species, it reduced the abundance and diversity of soil biota. We observed contrasting effects of tree‐ and soil‐related fire severity on survival and recovery of understory vegetation and soil biological communities. Severe fires that killed overstory Pinus sylvestris promoted a successional stage dominated by the mosses Ceratodon purpureus and Polytrichum juniperinum, but reduced regeneration of tree seedlings and disfavored the ericaceous dwarf‐shrub Vaccinium vitis‐idaea and the grass Deschampsia flexuosa. Moreover, high tree mortality from fire reduced fungal biomass and changed fungal community composition, in particular that of ectomycorrhizal fungi, and reduced the fungivorous soil Oribatida. In contrast, soil‐related fire severity had little impact on vegetation composition, fungal communities, and soil animals. Bacterial communities responded to both tree‐ and soil‐related fire severity. Synthesis: Our results 2 years postfire suggest that a change in fire regime from a historically low‐severity ground fire regime, with fires that mainly burns into the soil organic layer, to a stand‐replacing fire regime with a high degree of tree mortality, as may be expected with climate change, is likely to impact the short‐term recovery of stand structure and above‐ and belowground species composition of even‐aged P. sylvestris boreal forests.

Shrub abundance is expected to increase with enhanced temperature and nutrient availability in the Arctic, and associated changes in abundance of ectomycorrhizal (EM) fungi could be a key link between plant responses and longer-term changes in soil organic matter storage. This study quantifies the response in EM fungal abundance to long-term warming and fertilization in two arctic ecosystems with contrasting responses of the EM shrub Betula nana. Ergosterol was used as a biomarker for living fungal biomass in roots and organic soil and ingrowth bags were used to estimate EM mycelial production. We measured 15N and 13C natural abundance to identify the EM-saprotrophic divide in fungal sporocarps and to validate the EM origin of mycelia in the ingrowth bags. Fungal biomass in soil and EM mycelial production increased with fertilization at both tundra sites, and with warming at one site. This was caused partly by increased dominance of EM plants and partly by stimulation of EM mycelial growth. We conclude that cycling of carbon and nitrogen through EM fungi will increase when strongly nutrient-limited arctic ecosystems are exposed to a warmer and more nutrient-rich environment. This has potential consequences for below-ground litter quality and quantity, and for accumulation of organic matter in arctic soils.

Alterations in fire activity due to climate change and fire suppression may have profound effects on the balance between storage and release of carbon (C) and associated volatile elements. Stored soil mercury (Hg) is known to volatilize due to wildfires and this could substantially affect the land-air exchange of Hg; conversely the absence of fires and human disturbance may increase the time period over which Hg is sequestered. Here we show for a wildfire chronosequence spanning over more than 5000 years in boreal forest in northern Sweden that belowground inventories of total Hg are strongly related to soil humus C accumulation (R2 = 0.94, p < 0.001). Our data clearly show that northern boreal forest soils have a strong sink capacity for Hg, and indicate that the sequestered Hg is bound in soil organic matter pools accumulating over millennia. Our results also suggest that more than half of the Hg stock in the sites with the longest time since fire originates from deposition predating the onset of large-scale anthropogenic emissions. This study emphasizes the importance of boreal forest humus soils for Hg storage and reveals that this pool is likely to persist over millennial time scales in the prolonged absence of fire.

Summary 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.