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Enhanced nitrogen (N) availability in temperate forests has altered ecosystem carbon (C) and N cycling. Recent research has shown that these alterations lead to reductions in belowground C allocation by trees and that the consequences of these reductions on soil C and nutrient cycling may vary by mycorrhizal type. We hypothesized that trees that associate with ectomycorrhizal fungi (ECM) would reduce C allocation towards roots and mycorrhizal fungi to a greater extent than trees that associate with arbuscular mycorrhizal fungi (AM) in response to > 25 years of N fertilization. We further hypothesized that N induced decoupling of roots and microbes in ECM trees would be evidenced by greater declines in extracellular enzyme activities. We measured belowground C allocation to fine root biomass and mycorrhizal colonization in 6 AM and 6 ECM dominated plots in the N fertilized and reference watersheds at the Fernow Experimental Forest in West Virginia, USA. We compared these to measurements of simple-C, complex-C, nitrogen, and phosphorus acquiring enzyme activities in organic horizon, bulk mineral, and rhizosphere soil fractions. N fertilization reduced fine root biomass and mycorrhizal colonization in both AM and ECM stands. We found more consistent reductions in enzyme activities in ECM soils than AM soils under N fertilization which may have been driven by greater declines in root-C transfer to soil microbes. This mechanism helps to explain variability in soil C cycling responses across N gradient and fertilization experiments and may prove useful in predicting the fate of soil C stocks in response to N deposition. [ABSTRACT FROM AUTHOR]

With global change already enhancing drought frequency and severity, there is a critical need to determine if temperate forests will continue to act as carbon (C) sinks. The degree to which belowground strategies sustain tree growth under water stress remains uncertain. We examined differences between two temperate forest tree species in their belowground response to experimentally reduced precipitation over three years. We chose trees that are dominant in temperate forests and have different belowground resource acquisition strategies. We tested our hypothesis that in response to reduced precipitation, sugar maple trees would reduce belowground C investment while oak trees would show resiliency belowground. We found that oak trees maintained belowground C investment at the same level, likely reflecting an ability to leverage a higher baseline level of investment to enhance water and nutrient uptake. By contrast, sugar maple trees initially responded with increased investment in roots and mycorrhizae, but mycorrhizal colonization faded over time. Moreover, the treatment reduced rhizodeposition belowground by sugar maple trees, suggesting that the belowground C investment we observed does not lead to increased mobilization of nutrients. Our findings indicate that there are species specific responses of C investment in roots, mycorrhizae, and rhizodeposits to reduced precipitation. [ABSTRACT FROM AUTHOR]

Given that global change is predicted to increase the frequency and severity of drought in temperate forests, it is critical to understand the degree to which plant belowground responses cascade through the soil system to drive ecosystem responses to water stress. While most research has focused on plant and microbial responses independently of each other, a gap in our understanding lies in the integrated response of plant‐microbial interactions to water stress. We investigated the extent to which divergent belowground responses to reduced precipitation between sugar maple trees (Acer saccharum) versus oak trees (Oak spp.) may influence microbial activity via throughfall exclusion in the field. Evidence that oak trees send carbon belowground to prime microbial activity more than maples under ambient conditions and in response to water stress suggests there is the potential for corresponding impacts of reduced precipitation on microbial activity. As such, we tested the hypothesis that differences in belowground C allocation between oaks and maples would stimulate microbial activity in the oak treatment soils and reduce microbial activity in in the sugar maple treatment soils compared to their respective controls. We found that the treatment led to declines in N mineralization, soil respiration, and oxidative enzyme activity in the sugar maple treatment plot. These declines may be due to sugar maple trees reducing root C transfers to the soil. By contrast, the reduced precipitation treatment enhanced soil respiration, as well as rates of N mineralization and peroxidase activity in the oak rhizosphere. This enhanced activity suggests that oak roots provided optimal rhizosphere conditions during water stress to prime microbial activity to support net primary production. With future changes in precipitation predicted for forests in the Eastern US, we show that the strength of plant‐microbial interactions drives the degree to which reduced precipitation impacts soil C and nutrient cycling. [ABSTRACT FROM AUTHOR]

Conceptual and empirical advances in soil biogeochemistry have challenged long-held assumptions about the role of soil micro-organisms in soil organic carbon (SOC) dynamics; yet, rigorous tests of emerging concepts remain sparse. Recent hypotheses suggest that microbial necromass production links plant inputs to SOC accumulation, with high-quality (i.e., rapidly decomposing) plant litter promoting microbial carbon use efficiency, growth, and turnover leading to more mineral stabilization of necromass. We test this hypothesis experimentally and with observations across six eastern US forests, using stable isotopes to measure microbial traits and SOC dynamics. Here we show, in both studies, that microbial growth, efficiency, and turnover are negatively (not positively) related to mineral-associated SOC. In the experiment, stimulation of microbial growth by high-quality litter enhances SOC decomposition, offsetting the positive effect of litter quality on SOC stabilization. We suggest that microbial necromass production is not the primary driver of SOC persistence in temperate forests. Factors such as microbial necromass origin, alternative SOC formation pathways, priming effects, and soil abiotic properties can strongly decouple microbial growth, efficiency, and turnover from mineral-associated SOC. Mineral-associated soil carbon buildup is poorly explained by microbial necromass production (a common hypothesis). During litter decomposition, these processes are decoupled by priming effects and alternate soil carbon formation pathways [ABSTRACT FROM AUTHOR]

Summary: Decades of atmospheric nitrogen (N) deposition in the northeastern USA have enhanced this globally important forest carbon (C) sink by relieving N limitation. While many N fertilization experiments found increased forest C storage, the mechanisms driving this response at the ecosystem scale remain uncertain.Following the optimal allocation theory, augmented N availability may reduce belowground C investment by trees to roots and soil symbionts. To test this prediction and its implications on soil biogeochemistry, we constructed C and N budgets for a long‐term, whole‐watershed N fertilization study at the Fernow Experimental Forest, WV, USA.Nitrogen fertilization increased C storage by shifting C partitioning away from belowground components and towards aboveground woody biomass production. Fertilization also reduced the C cost of N acquisition, allowing for greater C sequestration in vegetation. Despite equal fine litter inputs, the C and N stocks and C : N ratio of the upper mineral soil were greater in the fertilized watershed, likely due to reduced decomposition of plant litter.By combining aboveground and belowground data at the watershed scale, this study demonstrates how plant C allocation responses to N additions may result in greater C storage in both vegetation and soil. [ABSTRACT FROM AUTHOR]

Roots promote the formation of slow‐cycling soil carbon (C), yet we have a limited understanding of the magnitude and controls on this flux. We hypothesised arbuscular mycorrhizal (AM)‐ and ectomycorrhizal (ECM)‐associated trees would exhibit differences in root‐derived C accumulation in the soil, and that much of this C would be transferred into mineral‐associated pools. We installed δ13C‐enriched ingrowth cores across mycorrhizal gradients in six Eastern U.S. forests (n = 54 plots). Overall, root‐derived C was 54% greater in AM versus ECM‐dominated plots. This resulted in nearly twice as much root‐derived C in putatively slow‐cycling mineral‐associated pools in AM compared to ECM plots. Given that our estimates of root‐derived inputs were often equal to or greater than leaf litter inputs, our results suggest that variation in root‐derived soil C accumulation due to tree mycorrhizal dominance may be a key control of soil C dynamics in forests. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Temperature and substrate availability constrain the activity of the extracellular enzymes that decompose and release nutrients from soil organic matter (SOM). Proteolytic enzymes are the primary class of enzymes involved in the depolymerization of nitrogen (N) from proteinaceous components of SOM, and their activity affects the rate of N cycling in forest soils. The objectives of this study were to determine whether and how temperature and substrate availability affect the activity of proteolytic enzymes in temperate forest soils, and whether the activity of proteolytic enzymes and other enzymes involved in the acquisition of N (i.e., chitinolytic and ligninolytic enzymes) differs between trees species that form associations with either ectomycorrhizal or arbuscular mycorrhizal fungi. Temperature limitation of proteolytic enzyme activity was observed only early in the growing season when soil temperatures in the field were near 4°C. Substrate limitation to proteolytic activity persisted well into the growing season. Ligninolytic enzyme activity was higher in soils dominated by ectomycorrhizal associated tree species. In contrast, the activity of proteolytic and chitinolytic enzymes did not differ, but there were differences between mycorrhizal association in the control of roots on enzyme activity. Roots of ectomycorrhizal species but not those of arbuscular mycorrhizal species exerted significant control over proteolytic, chitinolytic, and ligninolytic enzyme activity; the absence of ectomycorrhizal fine roots reduced the activity of all three enzymes. These results suggest that climate warming in the absence of increases in substrate availability may have a modest effect on soil-N cycling, and that global changes that alter below ground carbon allocation by trees are likely to have a larger effect on nitrogen cycling in stands dominated by ectomycorrhizal fungi. [ABSTRACT FROM AUTHOR]