Soil fungal and bacterial communities reflect differently tundra vegetation state transitions and soil physico-chemical properties.

Strong disturbances may induce ecosystem transitions into new alternative states that sustain through plant-soil interactions, such as the transition of dwarf shrub-dominated into graminoid-dominated vegetation by herbivory in tundra. Little evidence exists on soil microbial communities in alternative states, and along the slow process of ecosystem return into the predisturbance state. We analysed vegetation, soil microbial communities and activities as well as soil physico-chemical properties in historical reindeer enclosures in northernmost Finland in the following plot types: control heaths in the surrounding tundra; graminoid-dominated; 'shifting'; and recovered dwarf shrub-dominated vegetation inside enclosures. Soil fungal communities followed changes in vegetation, whereas bacterial communities were more affected by soil physico-chemical properties. Graminoid plots were characterized by moulds, pathotrophs and dark septate endophytes. Ericoid mycorrhizal and saprotrophic fungi were typical for control and recovered plots. Soil microbial communities inside the enclosures showed historical contingency, as their spatial variation was high in recovered plots despite the vegetation being more homogeneous. Self-maintaining feedback loops between plant functional types, soil microbial communities, and carbon and nutrient mineralization act effectively to stabilize alternative vegetation states, but once predisturbance vegetation reestablishes itself, soil microbial communities and physico-chemical properties return back towards their predisturbance state.

Ligneous amendments increase soil organic carbon content in fine-textured boreal soils and modulate N2O emissions.

Organic soil amendments are used to improve soil quality and mitigate climate change. However, their effects on soil structure, nutrient and water retention as well as greenhouse gas (GHG) emissions are still poorly understood. The purpose of this study was to determine the residual effects of a single field application of four ligneous soil amendments on soil structure and GHG emissions. We conducted a laboratory incubation experiment using soil samples collected from an ongoing soil-amendment field experiment at Qvidja Farm in south-west Finland, two years after a single application of four ligneous biomasses. Specifically, two biochars (willow and spruce) produced via slow pyrolysis, and two mixed pulp sludges from paper industry side-streams were applied at a rate of 9-22 Mg ha-1 mixed in the top 0.1 m soil layer. An unamended fertilized soil was used as a control. The laboratory incubation lasted for 33 days, during which the samples were kept at room temperature (21°C) and at 20%, 40%, 70% or 100% water holding capacity. Carbon dioxide (CO2), nitrous oxide (N2O) and methane (CH4) fluxes were measured periodically after 1, 5, 12, 20 and 33 days of incubation. The application of ligneous soil amendments increased the pH of the sampled soils by 0.4-0.8 units, whereas the effects on soil organic carbon content and soil structure varied between treatments. The GHG exchange was dominated by CO2 emissions, which were mainly unaffected by the soil amendment treatments. The contribution of soil CH4 exchange was negligible (nearly no emissions) compared to soil CO2 and N2O emissions. The soil N2O emissions exhibited a positive exponential relationship with soil moisture. Overall, the soil amendments reduced N2O emissions on average by 13%, 64%, 28%, and 37%, at the four soil moisture levels, respectively. Furthermore, the variation in N2O emissions between the amendments correlated positively with their liming effect. More specifically, the potential for the pulp sludge treatments to modulate N2O emissions was evident only in response to high water contents. This tendency to modulate N2O emissions was attributed to their capacity to increase soil pH and influence soil processes by persisting in the soil long after their application.

Moth outbreaks alter root-associated fungal communities in subarctic mountain birch forests.

Climate change has important implications on the abundance and range of insect pests in forest ecosystems. We studied responses of root-associated fungal communities to defoliation of mountain birch hosts by a massive geometrid moth outbreak through 454 pyrosequencing of tagged amplicons of the ITS2 rDNA region. We compared fungal diversity and community composition at three levels of moth defoliation (intact control, full defoliation in one season, full defoliation in two or more seasons), replicated in three localities. Defoliation caused dramatic shifts in functional and taxonomic community composition of root-associated fungi. Differentially defoliated mountain birch roots harbored distinct fungal communities, which correlated with increasing soil nutrients and decreasing amount of host trees with green foliar mass. Ectomycorrhizal fungi (EMF) abundance and richness declined by 70-80 % with increasing defoliation intensity, while saprotrophic and endophytic fungi seemed to benefit from defoliation. Moth herbivory also reduced dominance of Basidiomycota in the roots due to loss of basidiomycete EMF and increases in functionally unknown Ascomycota. Our results demonstrate the top-down control of belowground fungal communities by aboveground herbivory and suggest a marked reduction in the carbon flow from plants to soil fungi following defoliation. These results are among the first to provide evidence on cascading effects of natural herbivory on tree root-associated fungi at an ecosystem scale.

Short-term impacts of energy wood harvesting on ectomycorrhizal fungal communities of Norway spruce saplings.

The increased demand for harvesting energy wood raises questions about its effects on the functioning of the forest ecosystems, soil processes and biodiversity. Impacts of tree stump removal on ectomycorrhizal fungal (EMF) communities of Norway spruce saplings were studied with 454-pyrosequencing in a 3-year field experiment replicated in 3 geographical areas. This is possibly the most thorough investigation of EMF communities associated with saplings grown on sites subjected to energy wood harvesting. To separate impacts of tree stump and logging residue removal on EMF and plant variables, we used three harvesting treatments with increasing complexity from patch mounding alone (P) to patch mounding combined with logging residue removal (RP), and patch mounding combined with both logging residue and stump removal (SRP). Saplings grown in uncut forests (F) served as references for harvesting treatments. A majority of sequences (>92%) and operational taxonomic units (OTUs, 55%) were assigned as EMF. EMF OTU richness, fungal community composition or sapling growth did not differ between harvesting treatments (P, RP and SRP), while EMF OTU richness, diversity and evenness were highest and sapling growth lowest in the undisturbed reference forests (F). The short study period may partially explain the similarities in fungal and sapling variables in different harvesting treatments. In conclusion, our results indicate that neither stump removal nor logging residue removal have significant additional negative impacts on EMF communities or growth of Norway spruce saplings in the short-term compared with the impacts of more conventional harvesting methods, including clear cutting and patch mounding.

Moth herbivory enhances resource turnover in subarctic mountain birch forests?

Massive moth outbreaks cause large-scale damage in subarctic mountain birch forests with a concomitant decrease in carbon flux to mycorrhizal fungi and an increased deposition of dissolved carbon and nutrients as moth frass into soil. We investigated impacts of moth herbivory along three replicated gradients with three levels of moth herbivory (undamaged, once damaged, repeatedly damaged) on soil nutrient levels and biological parameters. We found an increase in soil nutrients and in the biomass of enchytraeid worms, which are key faunal decomposers. Fungi bacteria ratio and C:N ratio decreased in humus with increasing severity of herbivory. Our findings suggest enhanced resource turnover in mountain birch forests due to massive moth herbivory. This may provide a shortcut for carbon and nutrient input to subarctic soils, which largely bypasses the main routes of carbon from plants to soil via mycorrhizal and litter-decomposing fungi. Moreover, a temporal shift occurs in carbon allocation to soil, providing decomposers an opportunity to use an early-season peak in resource availability. Our results suggest a hitherto unappreciated role of massive insect herbivore attacks on resource dynamics in subarctic ecosystems.