Watershed-scale liming reveals the short- and long-term effects of pH on the forest soil microbiome and carbon cycling.

Soil microbial community composition routinely correlates with pH, reflecting both direct pH effects on microbial physiology and long-term biogeochemical feedbacks. We used two watershed-scale liming experiments to identify short- (2 years) and long-term (25 years) changes in the structure and function of bacterial and fungal communities in organic horizons (Oe and Oa ) of acid forest soils. Liming increased soil pH, extractable calcium, and soil carbon stocks, reduced biomass-specific respiration, and caused major changes in the soil microbiome in the short and long term. More taxa responded to liming in the short term (70%) than in the long term (30%), with most showing consistent directional responses at both sites. The ratio of change in relative abundance between limed and reference sites was twofold higher at the long than the short-term site, indicating that the effects of liming grew over time. Liming impacts were most pronounced in fungi, as steep declines of dominant ectomycorrhizal fungi (Cenococcum and Russula) occurred at both sites. Liming favoured neutrophilic bacteria over acidophilic populations according to estimated environmental pH optima. Collectively, these results demonstrate that a liming-induced change of one pH unit has an immediate and persistent effect on the structure and function of microbial communities in acid forest soils. The corresponding suppression of respiration indicates that anthropogenic alterations of soil pH, as driven by acid deposition or liming, can affect forest floor C stocks due to pH-driven shifts in community structure.

Microbial community shifts correspond with suppression of decomposition 25 years after liming of acidic forest soils.

Microbial community structure and function regularly covary with soil pH, yet effects of these interactions on soil carbon are rarely tested experimentally within natural ecosystems. We investigated the enduring (25 year) impacts of liming on microbial community structure and decomposition at an acidic northern hardwood forest, where experimental liming increased pH one unit and surprisingly doubled the organic carbon stocks of the forest floor. We show that this increase in carbon storage corresponded with restructuring of the bacterial and fungal communities that drive decomposition. In the Oe horizon, liming reduced the activities of five extracellular enzymes that mediate decomposition, while the Oa horizon showed an especially large (64%) reduction in the activity of a sixth, peroxidase, which is an oxidative enzyme central to lignocellulose degradation. Decreased enzyme activities corresponded with loss of microbial taxa important for lignocellulose decay, including large reductions in the dominant ectomycorrhizal genera Russula and Cenococcum, saprotrophic and wood decaying fungi, and Actinobacteria (Thermomonosporaceae). These results demonstrate the importance of pH as a dominant regulator of microbial community structure and illustrate how changes to this structure can produce large, otherwise unexpected increases in carbon storage in forest soils.

Unprocessed Atmospheric Nitrate in Waters of the Northern Forest Region in the U.S. and Canada.

Little is known about the regional extent and variability of nitrate from atmospheric deposition that is transported to streams without biological processing in forests. We measured water chemistry and isotopic tracers (δ18O and δ15N) of nitrate sources across the Northern Forest Region of the U.S. and Canada and reanalyzed data from other studies to determine when, where, and how unprocessed atmospheric nitrate was transported in catchments. These inputs were more widespread and numerous than commonly recognized, but with high spatial and temporal variability. Only 6 of 32 streams had high fractions (>20%) of unprocessed atmospheric nitrate during baseflow. Seventeen had high fractions during stormflow or snowmelt, which corresponded to large fractions in near-surface soil waters or groundwaters, but not deep groundwater. The remaining 10 streams occasionally had some (<20%) unprocessed atmospheric nitrate during stormflow or baseflow. Large, sporadic events may continue to be cryptic due to atmospheric deposition variation among storms and a near complete lack of monitoring for these events. A general lack of observance may bias perceptions of occurrence; sustained monitoring of chronic nitrogen pollution effects on forests with nitrate source apportionments may offer insights needed to advance the science as well as assess regulatory and management schemes.

Effects of climate warming on carbon fluxes in grasslands- A global meta-analysis.

Climate warming will affect terrestrial ecosystems in many ways, and warming-induced changes in terrestrial carbon (C) cycling could accelerate or slow future warming. So far, warming experiments have shown a wide range of C flux responses, across and within biome types. However, past meta-analyses of C flux responses have lacked sufficient sample size to discern relative responses for a given biome type. For instance grasslands contribute greatly to global terrestrial C fluxes, and to date grassland warming experiments provide the opportunity to evaluate concurrent responses of both plant and soil C fluxes. Here, we compiled data from 70 sites (in total 622 observations) to evaluate the response of C fluxes to experimental warming across three grassland types (cold, temperate, and semi-arid), warming methods, and short (≤3 years) and longer-term (>3 years) experiment lengths. Overall, our meta-analysis revealed that experimental warming stimulated C fluxes in grassland ecosystems with regard to both plant production (e.g., net primary productivity (NPP) 15.4%; aboveground NPP (ANPP) by 7.6%, belowground NPP (BNPP) by 11.6%) and soil respiration (Rs) (9.5%). However, the magnitude of C flux stimulation varied significantly across cold, temperate and semi-arid grasslands, in that responses for most C fluxes were larger in cold than temperate or semi-arid ecosystems. In semi-arid and temperate grasslands, ecosystem respiration (Reco) was more sensitive to warming than gross primary productivity (GPP), while the opposite was observed for cold grasslands, where warming produced a net increase in whole-ecosystem C storage. However, the stimulatory effect of warming on ANPP and Rs observed in short-term studies (≤3 years) in both cold and temperate grasslands disappeared in longer-term experiments (>3 years). These results highlight the importance of conducting long-term warming experiments, and in examining responses across a wide range of climate.

Beyond Static Benchmarking: Using Experimental Manipulations to Evaluate Land Model Assumptions.

Land models are often used to simulate terrestrial responses to future environmental changes, but these models are not commonly evaluated with data from experimental manipulations. Results from experimental manipulations can identify and evaluate model assumptions that are consistent with appropriate ecosystem responses to future environmental change. We conducted simulations using three coupled carbon-nitrogen versions of the Community Land Model (CLM, versions 4, 4.5, and-the newly developed-5), and compared the simulated response to nitrogen (N) and atmospheric carbon dioxide (CO2) enrichment with meta-analyses of observations from similar experimental manipulations. In control simulations, successive versions of CLM showed a poleward increase in gross primary productivity and an overall bias reduction, compared to FLUXNET-MTE observations. Simulations with N and CO2 enrichment demonstrate that CLM transitioned from a model that exhibited strong nitrogen limitation of the terrestrial carbon cycle (CLM4) to a model that showed greater responsiveness to elevated concentrations of CO2 in the atmosphere (CLM5). Overall, CLM5 simulations showed better agreement with observed ecosystem responses to experimental N and CO2 enrichment than previous versions of the model. These simulations also exposed shortcomings in structural assumptions and parameterizations. Specifically, no version of CLM captures changes in plant physiology, allocation, and nutrient uptake that are likely important aspects of terrestrial ecosystems' responses to environmental change. These highlight priority areas that should be addressed in future model developments. Moving forward, incorporating results from experimental manipulations into model benchmarking tools that are used to evaluate model performance will help increase confidence in terrestrial carbon cycle projections.

Multiyear fate of a 15 N tracer in a mixed deciduous forest: retention, redistribution, and differences by mycorrhizal association.

The impact of atmospheric nitrogen deposition on forest ecosystems depends in large part on its fate. Past tracer studies show that litter and soils dominate the short-term fate of added 15 N, yet few have examined its longer term dynamics or differences among forest types. This study examined the fate of a 15 N-NO3- tracer over 5-6 years in a mixed deciduous stand that was evenly composed of trees with ectomycorrhizal and arbuscular mycorrhizal associations. The tracer was expected to slowly mineralize from its main initial fate in litter and surface soil, with some 15 N moving to trees, some to deeper soil, and some net losses. Recovery of added 15 N in trees and litterfall totaled 11.3% both 1 and 5-6 years after the tracer addition, as 15 N redistributed from fine and especially coarse roots into cumulative litterfall and small accumulations in woody tissues. Estimates of potential carbon sequestration from tree 15 N recovery amounted to 12-14 kg C per kg of N deposition. Tree 15 N acquisition occurred within the first year after the tracer addition, with no subsequent additional net transfer of 15 N from detrital to plant pools. In both years, ectomycorrhizal trees gained 50% more of the tracer than did trees with arbuscular mycorrhizae. Much of the 15 N recovered in wood occurred in tree rings formed prior to the 15 N addition, demonstrating the mobility of N in wood. Tracer recovery rapidly decreased over time in surface litter material and accumulated in both shallow and deep soil, perhaps through mixing by earthworms. Overall, results showed redistribution of tracer 15 N through trees and surface soils without any losses, as whole-ecosystem recovery remained constant between 1 and 5-6 years at 70% of the 15 N addition. These results demonstrate the persistent ecosystem retention of N deposition even as it redistributes, without additional plant uptake over this timescale.

Soil processes drive seasonal variation in retention of 15N tracers in a deciduous forest catchment.

Seasonal patterns of stream nitrate concentration have long been interpreted as demonstrating the central role of plant uptake in regulating stream nitrogen loss from forested catchments. Soil processes are rarely considered as important drivers of these patterns. We examined seasonal variation in N retention in a deciduous forest using three whole-ecosystem 15N tracer additions: in late April (post-snowmelt, pre-leaf-out), late July (mid-growing- season), and late October (end of leaf-fall). We expected that plant 15N uptake would peak in late spring and midsummer, that immobilization in surface litter and soil would peak the following autumn leaf-fall, and that leaching losses would vary inversely with 15N retention. Similar to most other 15N tracer studies, we found that litter and soils dominated ecosystem retention of added 15N. However, 15N recovery in detrital pools varied tremendously by season, with > 90% retention in spring and autumn and sharply reduced 15N retention in late summer. During spring, over half of the 15N retained in soil occurred within one day in the heavy (mineral-associated) soil fraction. During summer, a large decrease in 15N retention one week after addition coincided with increased losses of 15NO3- to soil leachate and seasonal increases in soil and stream NO3- concentrations, although leaching accounted for only a small fraction of the lost 15N (< 0.2%). Uptake of 15N into roots did not vary by season and accounted for < 4% of each tracer addition. Denitrification or other processes that lead to N gas loss may have consumed the rest. These measurements of 15N movement provide strong evidence for the dominant role of soil processes in regulating seasonal N retention and losses in this catchment and perhaps others with similar soils.

Isotopic signals of summer denitrification in a northern hardwood forested catchment.

Despite decades of measurements, the nitrogen balance of temperate forest catchments remains poorly understood. Atmospheric nitrogen deposition often greatly exceeds streamwater nitrogen losses; the fate of the remaining nitrogen is highly uncertain. Gaseous losses of nitrogen to denitrification are especially poorly documented and are often ignored. Here, we provide isotopic evidence (δ(15)NNO3 and δ(18)ONO3) from shallow groundwater at the Hubbard Brook Experimental Forest indicating extensive denitrification during midsummer, when transient, perched patches of saturation developed in hillslopes, with poor hydrological connectivity to the stream, while streamwater showed no isotopic evidence of denitrification. During small rain events, precipitation directly contributed up to 34% of streamwater nitrate, which was otherwise produced by nitrification. Together, these measurements reveal the importance of denitrification in hydrologically disconnected patches of shallow groundwater during midsummer as largely overlooked control points for nitrogen loss from temperate forest catchments.

Global patterns of nitrogen limitation: confronting two global biogeochemical models with observations.

Projections of future changes in land carbon (C) storage using biogeochemical models depend on accurately modeling the interactions between the C and nitrogen (N) cycles. Here, we present a framework for analyzing N limitation in global biogeochemical models to explore how C-N interactions of current models compare to field observations, identify the processes causing model divergence, and identify future observation and experiment needs. We used a set of N-fertilization simulations from two global biogeochemical models (CLM-CN and O-CN) that use different approaches to modeling C-N interactions. On the global scale, net primary productivity (NPP) in the CLM-CN model was substantially more responsive to N fertilization than in the O-CN model. The most striking difference between the two models occurred for humid tropical forests, where the CLM-CN simulated a 62% increase in NPP at high N addition levels (30 g N m(-2) yr(-1)), while the O-CN predicted a 2% decrease in NPP due to N fertilization increasing plant respiration more than photosynthesis. Across 35 temperate and boreal forest sites with field N-fertilization experiments, we show that the CLM-CN simulated a 46% increase in aboveground NPP in response to N, which exceeded the observed increase of 25%. In contrast, the O-CN only simulated a 6% increase in aboveground NPP at the N-fertilization sites. Despite the small response of NPP to N fertilization, the O-CN model accurately simulated ecosystem retention of N and the fate of added N to vegetation when compared to empirical (15) N tracer application studies. In contrast, the CLM-CN predicted lower total ecosystem N retention and partitioned more losses to volatilization than estimated from observed N budgets of small catchments. These results point to the need for model improvements in both models in order to enhance the accuracy with which global C-N cycle feedbacks are simulated.

Forest liming increases forest floor carbon and nitrogen stocks in a mixed hardwood forest.

In acid-impacted forests, decreased soil pH and calcium (Ca) availability have the potential to influence biotic and abiotic controls on carbon (C) and nitrogen (N) cycling. We investigated the effects of liming on above- and belowground C and N pools and fluxes 19 years after lime addition to the Woods Lake Watershed, Adirondack Park, New York, USA. Soil pH and exchangeable Ca remained elevated in the forest floor and upper mineral soil of limed areas. Forest floor C and N stocks were significantly larger in limed plots (68 vs. 31 Mg C/ha, and 3.0 vs. 1.5 Mg N/ha), resulting from a larger mass of Oa material. Liming reduced soil basal respiration rates by 17% and 43% in the Oe and Oa horizons, respectively. Net N mineralization was significantly lower in the limed soils for both forest floor horizons. Additional measurements of forest floor depth outside of our study plots, but within the treatment and control subcatchments also showed a deeper forest floor in limed areas; however, the mean depth of limed forest floor was 5 cm shallower than that observed in our study plots. Using a differential equation model of forest floor C dynamics, we found that liming effects on C fluxes measured within our study plots could explain the small observed increase in the Oe C stock but were not large enough to explain the increase in the Oa. Our catchment-wide assessment of forest floor depth, however, indicates that our plot analysis may be an overestimate of ecosystem-scale C and N stocks. Our results suggest that the mechanisms identified in our study, primarily liming-induced reduction in decomposition rates, may account for much of the observed increase in forest floor C. These findings emphasize the importance of understanding of the effects of liming in hardwood forests, and the long-term impacts of acid deposition on forest C and N uptake and retention.

Climate change impacts of US reactive nitrogen.

Fossil fuel combustion and fertilizer application in the United States have substantially altered the nitrogen cycle, with serious effects on climate change. The climate effects can be short-lived, by impacting the chemistry of the atmosphere, or long-lived, by altering ecosystem greenhouse gas fluxes. Here we develop a coherent framework for assessing the climate change impacts of US reactive nitrogen emissions, including oxides of nitrogen, ammonia, and nitrous oxide (N(2)O). We use the global temperature potential (GTP), calculated at 20 and 100 y, in units of CO(2) equivalents (CO(2)e), as a common metric. The largest cooling effects are due to combustion sources of oxides of nitrogen altering tropospheric ozone and methane concentrations and enhancing carbon sequestration in forests. The combined cooling effects are estimated at -290 to -510 Tg CO(2)e on a GTP(20) basis. However, these effects are largely short-lived. On a GTP(100) basis, combustion contributes just -16 to -95 Tg CO(2)e. Agriculture contributes to warming on both the 20-y and 100-y timescales, primarily through N(2)O emissions from soils. Under current conditions, these warming and cooling effects partially offset each other. However, recent trends show decreasing emissions from combustion sources. To prevent warming from US reactive nitrogen, reductions in agricultural N(2)O emissions are needed. Substantial progress toward this goal is possible using current technology. Without such actions, even greater CO(2) emission reductions will be required to avoid dangerous climate change.