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Forest ecosystems are subjected to global change drivers worldwide, such as increasing temperature, atmospheric carbon dioxide, nutrient pollution, as well as changes in fire and precipitation regimes. These global change drivers have greatly modified the biogeochemical cycles of carbon (C), nitrogen (N), and phosphorus (P), which has an impact on primary productivity in forest ecosystems and in turn, affect the quality and quantity of resources entering the soil food web. However, C, N, and P soil dynamics have been mostly studied without considering their coupling effects on soil organisms. This is of critical interest because changes in nutrient stoichiometry may have a strong effect on soil biota and the ecosystem functions they drive. Further, most studies have focused on global change effects on bacteria and fungi and their C:N:P stoichiometry, while neglecting other soil organisms at higher trophic levels. This has led to an incomplete understanding of how the entire soil food web drives ecosystem processes involved in organic matter turnover and nutrient cycling. Here, we review studies that investigated how global change drivers impact C:N:P stoichiometry of soil organisms at different trophic levels in forest ecosystems and identify important knowledge gaps. We propose future directions for research on global change impacts on the linkages between soil biota and C:N:P stoichiometry.

There is evidence that anthropogenic nitrogen (N) deposition enhances carbon (C) sequestration in boreal forest soils. However, it is unclear how free-living saprotrophs (bacteria and fungi, SAP) and ectomycorrhizal (EM) fungi responses to N addition impact soil C dynamics. Our aim was to investigate how SAP and EM communities are impacted by N enrichment and to estimate whether these changes influence decay of litter and humus. We conducted a long-term experiment in northern Sweden, maintained since 2004, consisting of ambient, low N additions (0, 3, 6, and 12 kg N ha(-1) year(-1)) simulating current N deposition rates in the boreal region, as well as a high N addition (50 kg N ha(-1) year(-1)). Our data showed that long-term N enrichment impeded mass loss of litter, but not of humus, and only in response to the highest N addition treatment. Furthermore, our data showed that EM fungi reduced the mass of N and P in both substrates during the incubation period compared to when only SAP organisms were present. Low N additions had no effect on microbial community structure, while the high N addition decreased fungal and bacterial biomasses and altered EM fungi and SAP community composition. Actinomycetes were the only bacterial SAP to show increased biomass in response to the highest N addition. These results provide a mechanistic understanding of how anthropogenic N enrichment can influence soil C accumulation rates and suggest that current N deposition rates in the boreal region (<= 12 kg N ha(-1) year(-1)) are likely to have a minor impact on the soil microbial community and the decomposition of humus and litter.

Nutrient optimization has been proposed as a way to increase boreal forest production, and involves chronic additions of liquid fertilizer with amounts of micro- and macro-nutrients adjusted annually to match tree nutritional requirements. We used a short-term (maintained since 2007) and a long-term (maintained since 1987) fertilization experiment in northern Sweden, in order to understand nutrient optimization effects on soil microbiota and mesofauna, and to explore the relationships between plant litter and microbial elemental stoichiometry. Soil microbes, soil fauna, and aboveground litter were collected from the control plots, and short- and long-term nutrient optimization plots. Correlation analyses revealed no relationships between microbial biomass and litter nutrient ratios. Litter C:N, C:P and N:P ratios declined in response to both optimization treatments; while only microbial C:P ratios declined in response to long-term nutrient optimization. Further, we found that both short- and long-term optimization treatments decreased total microbial, fungal, and bacterial PLFA biomass and shifted the microbial community structure towards a lower fungi:bacterial ratio. In contrast, abundances of most fungal- and bacterial-feeding soil biota were little affected by the nutrient optimization treatments. However, abundance of hemi-edaphic Collembola declined in response to the long-term nutrient optimization treatment. The relative abundances (%) of fungal-feeding and plant-feeding nematodes, respectively, declined and increased in response to both short-term and long-term treatments; bacterial-feeding nematodes increased relative to fungal feeders. Overall, our results demonstrate that long-term nutrient optimization aiming to increase forest production decreases litter C:N, C:P and N:P ratios, microbial C:P ratios and fungal biomass, whereas higher trophic levels are less affected.

There is evidence that anthropogenic nitrogen (N) deposition enhances carbon (C) sequestration in boreal soils. However, key underlying mechanisms explaining this increase have not been resolved. Two potentially important mechanisms are that aboveground litter production increases, or that litter quality changes in response to N enrichment. As such, our aim was to quantify whether simulated chronic N deposition caused changes in aboveground litter production or quality in a boreal forest. We conducted a long-term (17 years) stand-scale (0.1 ha) forest experiment, consisting of three N addition levels (0, 12.5, and 50 kg N ha(-1) yr(-1)) in northern Sweden, where background N deposition rates are very low. We measured the annual quantity of litter produced for 8 different litter categories, as well as their concentrations of C, N, phosphorus (P), lignin, cellulose and hemi-cellulose. Our results indicate that mosses were the only major litter component showing significant quantitative and qualitative alterations in response to the N additions, indicative of their ability to intercept a substantial portion of the N added. These effects were, however, offset by the other litter fractions where we found no changes in the total litter fluxes, or individual chemical constituents when all litter categories were summed. This study indicates that the current annual litter fluxes cannot explain the increase in soil C that has occurred in our study system in response to simulated chronic N application. These results suggest that other mechanisms are likely to explain the increased soil C accumulation rate we have observed, such as changes in soil microbial activity, or potentially transient changes in aboveground litter inputs that were no longer present at the time of our study.

It is proposed that carbon (C) sequestration in response to reactive nitrogen (N-r) deposition in boreal forests accounts for a large portion of the terrestrial sink for anthropogenic CO2 emissions. While studies have helped clarify the magnitude by which N-r deposition enhances C sequestration by forest vegetation, there remains a paucity of long-term experimental studies evaluating how soil C pools respond. We conducted a long-term experiment, maintained since 1996, consisting of three N addition levels (0, 12.5, and 50kgNha(-1)yr(-1)) in the boreal zone of northern Sweden to understand how atmospheric N-r deposition affects soil C accumulation, soil microbial communities, and soil respiration. We hypothesized that soil C sequestration will increase, and soil microbial biomass and soil respiration will decrease, with disproportionately large changes expected compared to low levels of N addition. Our data showed that the low N addition treatment caused a non-significant increase in the organic horizon C pool of similar to 15% and a significant increase of similar to 30% in response to the high N treatment relative to the control. The relationship between C sequestration and N addition in the organic horizon was linear, with a slope of 10kgCkg(-1)N. We also found a concomitant decrease in total microbial and fungal biomasses and a similar to 11% reduction in soil respiration in response to the high N treatment. Our data complement previous data from the same study system describing aboveground C sequestration, indicating a total ecosystem sequestration rate of 26kgCkg(-1)N. These estimates are far lower than suggested by some previous modeling studies, and thus will help improve and validate current modeling efforts aimed at separating the effect of multiple global change factors on the C balance of the boreal region.