Pulse additions of soil carbon and nitrogen affect soil nitrogen dynamics in an arid Colorado Plateau shrubland

Biogeochemical cycles in arid and semi-arid ecosystems depend upon the ability of soil microbes to use pulses of resources. Brief periods of high activity generally occur after precipitation events that provide access to energy and nutrients (carbon and nitrogen) for soil organisms. To better understand pulse-driven dynamics of microbial soil nitrogen (N) cycling in an arid Colorado Plateau ecosystem, we simulated a pulsed addition of labile carbon (C) and N in the field under the canopies of the major plant species in plant interspaces. Soil microbial activity and N cycling responded positively to added C while NH₄⁺–N additions resulted in an accumulation of soil NO₃⁻. Increases in microbial activity were reflected in higher rates of respiration and N immobilization with C addition. When both C and N were added to soils, N losses via NH₃ volatilization decreased. There was no effect of soil C or N availability on microbial biomass N suggesting that the level of microbial activity (respiration) may be more important than population size (biomass) in controlling short-term dynamics of inorganic and labile organic N. The effects of C and N pulses on soil microbial function and pools of NH₄⁺–N and labile organic N were observed to last only for the duration of the moisture pulse created by treatment addition, while the effect on the NO₃⁻–N pool persisted after soils dried to pre-pulse moisture levels. We observed that increases in available C lead to greater ecosystem immobilization and retention of N in soil microbial biomass and also lowered rates of gaseous N loss. With the exception of trace gas N losses, the lack of interaction between available C and N on controlling N dynamics, and the subsequent reduction in plant available N with C addition has implications for the competitive relationships between plants species, plants and microbes, or both.     

Soil nitrogen transformations under Populus tremuloides, Betula papyrifera and Acer saccharum following 3 years exposure to elevated CO2 and O3

Increases in atmospheric CO₂ and tropospheric O₃ may affect forest N cycling by altering plant litter production and the availability of substrates for microbial metabolism. Three years following the establishment of our free‐air CO₂–O₃ enrichment experiment, plant growth has been stimulated by elevated CO₂ resulting in greater substrate input to soil; elevated O₃ has counteracted this effect. We hypothesized that rates of soil N cycling would be enhanced by greater plant productivity under elevated CO₂, and that CO₂ effects would be dampened by O₃. We found that elevated CO₂ did not alter gross N transformation rates. Elevated O₃ significantly reduced gross N mineralization and microbial biomass N, and effects were consistent among species. We also observed significant interactions between CO₂ and O₃: (i) gross N mineralization was greater under elevated CO₂ (1.0 mg N kg^?1 day^?1) than in the presence of both CO₂ and O₃ (0.5 mg N kg^?1 day^?1) and (ii) gross NH₄⁺ immobilization was also greater under elevated CO₂ (0.8 mg N kg^?1 day^?1) than under CO₂ plus O₃ (0.4 mg N kg^?1 day^?1). We used a laboratory ¹⁵N tracer method to quantify transfer of inorganic N to organic pools. Elevated CO₂ led to greater recovery of NH₄⁺‐¹⁵N in microbial biomass and corresponding lower recovery in the extractable NO₃^? pool. Elevated CO₂ resulted in a substantial increase in NO₃^?‐¹⁵N recovery in soil organic matter. We observed no O₃ main effect and no CO₂ by O₃ interaction effect on ¹⁵N recovery in any soil pool. All of the above responses were most pronounced beneath Betula papyrifera and Populus tremuloides, which have grown more rapidly than Acer saccharum. Although elevated CO₂ has increased plant productivity, the resulting increase in plant litter production has yet to overcome the influence of the pre‐existing pool of soil organic matter on soil microbial activity and rates of N cycling. Ozone reduces plant litter inputs and also appears to affect the composition of plant litter in a way that reduces microbial biomass and activity.     

A meta‐analysis of soil salinization effects on nitrogen pools,cycles and fluxes in coastal ecosystems

Salinity intrusion caused by land subsidence resulting from increasing groundwater abstraction, decreasing river sediment loads and increasing sea level because of climate change has caused widespread soil salinization in coastal ecosystems. Soil salinization may greatly alter nitrogen (N) cycling in coastal ecosystems. However, a comprehensive understanding of the effects of soil salinization on ecosystem N pools, cycling processes and fluxes is not available for coastal ecosystems. Therefore, we compiled data from 551 observations from 21 peer‐reviewed papers and conducted a meta‐analysis of experimental soil salinization effects on 19 variables related to N pools, cycling processes and fluxes in coastal ecosystems. Our results showed that the effects of soil salinization varied across different ecosystem types and salinity levels. Soil salinization increased plant N content (18%), soil NH₄⁺ (12%) and soil total N (210%), although it decreased soil NO₃^? (2%) and soil microbial biomass N (74%). Increasing soil salinity stimulated soil N₂O fluxes as well as hydrological NH₄⁺ and NO₂^? fluxes more than threefold, although it decreased the hydrological dissolved organic nitrogen (DON) flux (59%). Soil salinization also increased the net N mineralization by 70%, although salinization effects were not observed on the net nitrification, denitrification and dissimilatory nitrate reduction to ammonium in this meta‐analysis. Overall, this meta‐analysis improves our understanding of the responses of ecosystem N cycling to soil salinization, identifies knowledge gaps and highlights the urgent need for studies on the effects of soil salinization on coastal agro‐ecosystem and microbial N immobilization. Additional increases in knowledge are critical for designing sustainable adaptation measures to the predicted intrusion of salinity intrusion so that the productivity of coastal agro‐ecosystems can be maintained or improved and the N losses and pollution of the natural environment can be minimized.     

Feedback of grazing on gross rates of N mineralization and inorganic N partitioning in steppe soils of Inner Mongolia

Plant-microbe interactions are crucial regulators of belowground nitrogen cycling in terrestrial ecosystems. However, such interactions have mostly been excluded from experimental setups for the investigation of gross inorganic N fluxes and N partitioning to plants and microorganisms. Ungulate grazing is likely to feed back on soil N fluxes, and hence it is of special importance to simultaneously investigate grazing effects on both plant and microbial N fluxes in intact plant-soil systems, where plant-microbe interactions persist during the experimental incubation. Based on the homogenous ¹⁵NH ₄ ⁺ labelling of intact plant-soil monoliths we investigated how various stocking rates (0, 2.35, 4.8 and 7.85 sheep ha^?1 grazing season^?1) in steppe of Inner Mongolia feedback on gross rates of N mineralization and short-term inorganic N partitioning between plant, microbial and soil N pools. Our results showed that the effect of grazing on gross N mineralization was non-uniform. At low stocking rate gross N mineralization tended to decrease but increased with higher grazing pressure. Hence, there was no significant correlation between stocking rate and gross N mineralization across the investigated grazing intensities. Grazing decreased ¹⁵N recovery both in plant and microbial N pools but strongly promoted NO ₃ ^? accumulation in the soil and thus negatively affected potential ecosystem N retention. This appeared to be closely related to the grazing-induced decline in easily degradable soil C availability at increasing stocking rate.     

Biotic and abiotic immobilization of ammonium, nitrite, and nitrate in soils developed under different tree species in the Catskill Mountains, New York, USA

Nitrogen retention in soil organic matter (SOM) is a key process influencing the accumulation and loss of N in forest ecosystems, but the rates and mechanisms of inorganic N retention in soils are not well understood. The primary objectives of this study were to compare ammonium (NH₄⁺), nitrite (NO₂^?), and nitrate (NO₃^?) immobilization among soils developed under different tree species in the Catskill Mountains of New York State, and to determine the relative roles of biotic or abiotic processes in soil N retention. A laboratory experiment was performed, where ¹⁵N was added as NH₄⁺, NO₂^?, or NO₃^? to live and mercury‐treated O horizon soils from three tree species (American beech, northern red oak, sugar maple), and ¹⁵N recoveries were determined in the SOM pool. Mercuric chloride was used to treat soils as this chemical inhibits microbial metabolism without significantly altering the chemistry of SOM. The recovery of ¹⁵N in SOM was almost always greater for NH₄⁺ (mean 20%) and NO₂^? (47%) than for NO₃^? (10%). Ammonium immobilization occurred primarily by biotic processes, with mean recoveries in live soils increasing from 9% at 15 min to 53% after 28 days of incubation. The incorporation of NO₂^? into SOM occurred rapidly (<15 min) via abiotic processes. Abiotic immobilization of NO₂^? (mean recovery 58%) was significantly greater than abiotic immobilization of NH₄⁺ (7%) or NO₃^? (7%). The incorporation of NO₂^? into SOM did not vary significantly among tree species, so this mechanism likely does not contribute to differences in soil NO₃^? dynamics among species. As over 30% of the ¹⁵NO₂^? label was recovered in SOM within 15 min in live soils, and the products of NO₂^? incorporation into SOM remained relatively stable throughout the 28‐day incubation, our results suggest that NO₂^? incorporation into SOM may be an important mechanism of N retention in forest soils. The importance of NO₂^? immobilization for N retention in field soils, however, will depend on the competition between incorporation into SOM and nitrification for transiently available NO₂^?. Further research is required to determine the importance of this process in field environments.     

Changes in nitrogen cycling and retention processes in soils under spruce forests along a nitrogen enrichment gradient in Germany

A network of long-term monitoring sites on nitrogen (N) input and output of forests across Germany showed that a number of Germany's forests are subject to or are experiencing N saturation and that spruce (Picea abies) stands have high risk. Our study was aimed at (1) quantifying the changes in gross rates of microbial N cycling and retention processes in forest soils along an N enrichment gradient and (2) relating the changes in soil N dynamics to N losses. We selected spruce sites representing an N enrichment gradient (indicated by leaching : throughfall N ratios) ranging from 0.04–0.13 (low N),≤0.26 (intermediate N enrichment) to≥0.42 (highly N enriched). To our knowledge, our study is the first to report on mechanistic changes in gross rates of soil N cycling and abiotic NO₃⁻ retention under ambient N enrichment gradient. Gross N mineralization, NH₄⁺ immobilization, gross nitrification, and NO₃⁻ immobilization rates increased up to intermediate N enrichment level and somewhat decreased at highly N-enriched condition. The turnover rates of NH₄⁺ and microbial N pools increased while the turnover rates of the NO₃⁻ pool decreased across the N enrichment gradient. Abiotic immobilization of NH₄⁺ did not differ across sites and was lower than that of NO₃⁻. Abiotic NO₃⁻ immobilization decreased across the N enrichment gradient. Microbial assimilation and turnover appeared to contribute largely to the retention of NH₄⁺. The increasing NO₃⁻ deposition and decreasing turnover rates of the NO₃⁻ pool, combined with decreasing abiotic NO₃⁻ retention, possibly contributed to increasing NO₃⁻ leaching and gaseous emissions across the N enrichment gradient. The empirical relationships of changes in microbial N cycling across the N enrichment gradient may be integrated in models used to predict responses of forest ecosystems (e.g. spruce) to increasing N deposition.     

Effects of plant species diversity, atmospheric [CO2], and N addition on gross rates of inorganic N release from soil organic matter

A significant challenge in predicting terrestrial ecosystem response to global changes comes from the relatively poor understanding of the processes that control pools and fluxes of plant nutrients in soil. In addition, individual global changes are often studied in isolation, despite the potential for interactive effects among them on ecosystem processes. We studied the response of gross N mineralization and microbial respiration after 6 years of application of three global change factors in a grassland field experiment in central Minnesota (the BioCON experiment). BioCON is a factorial manipulation of plant species diversity (1, 4, 9 and 16 prairie species), atmospheric [CO₂] (ambient and elevated: 560 μmol mol^?1), and N inputs (ambient and ambient +4 g N m^?2 yr^?1). We hypothesized that gross N mineralization would increase with increasing levels of all factors because of stimulated plant productivity and thus greater organic inputs to soils. However, we also hypothesized that N addition would enhance, while elevated [CO₂] and greater diversity would temper, gross N mineralization responses because of increased and reduced plant tissue N concentrations, respectively. In partial support of our hypothesis, gross N mineralization increased with greater diversity and N addition, but not with elevated [CO₂]. The ratio of gross N mineralization to microbial respiration (i.e. the ‘yield’ of inorganic N mineralized per unit C respired) declined with greater diversity and [CO₂] suggesting increasing limitation of microbial processes by N relative to C in these treatments. Based on these results, we conclude that the plant supply of organic matter primarily controls gross N mineralization and microbial respiration, but that the concentration of N in organic matter input secondarily influences these processes. Thus, in systems where N limits plant productivity these global change factors could cause different long‐term ecosystem trajectories because of divergent effects on soil N and C cycling.     

Disentangling species and functional group richness effects on soil N cycling in a grassland ecosystem

Species richness (SR) and functional group richness (FGR) are often confounded in both observational and experimental field studies of biodiversity and ecosystem function. This precludes discernment of their separate influences on ecosystem processes, including nitrogen (N) cycling, and how those influences might be moderated by global change factors. In a 17‐year field study of grassland species, we used two full factorial experiments to independently vary SR (one or four species, with FGR = 1) and FGR (1–4 groups, with SR = 4) to assess SR and FGR effects on ecosystem N cycling and its response to elevated carbon dioxide (CO₂) and N addition. We hypothesized that increased plant diversity (either SR or FGR) and elevated CO₂ would enhance plant N pools because of greater plant N uptake, but decrease soil N cycling rates because of greater soil carbon inputs and microbial N immobilization. In partial support of these hypotheses, increasing SR or FGR (holding the other constant) enhanced total plant N pools and decreased soil nitrate pools, largely through higher root biomass, and increasing FGR strongly reduced mineralization rates, because of lower root N concentrations. In contrast, increasing SR (holding FGR constant and despite increasing total plant C and N pools) did not alter root N concentrations or net N mineralization rates. Elevated CO₂ had minimal effects on plant and soil N metrics and their responses to plant diversity, whereas enriched N increased plant and soil N pools, but not soil N fluxes. These results show that functional diversity had additional effects on both plant N pools and rates of soil N cycling that were independent of those of species richness.     

Effect of Eucalyptus saligna and Albizia falcataria on soil processes and nitrogen supply in Hawaii

This study investigated the differences between two fast-growing tropical tree species on soil N flux and availability. The work was conducted in the island of Hawaii and included three sites located along the Hamakua coast on the northeastern side of the island. Within each site pure stands of Eucalyptus saligna (Sm.)␣and the N2-fixing Albizia falcataria (L.) Fosberg [=Paraserianthes falcataria (L.) Nielsen] were arranged in four randomized complete blocks. For most of the variables considered in this study, the species effects were usually strong and the site effects were significant in some cases. After 13 years, soils under the Albizia stand contained larger pools of total soil C and N, and larger pools of inorganic N. Soil N availability indexed by ion exchange resin bags revealed a strong pattern of species and site effect on N availability; soils under Albizia showed a 2.6–9 fold increase in N availability (P < 0.01). Potential net rates of N transformation (10- and 30-day aerobic incubations) were more than twice as high for soils under the Albizia than under the Eucalytus stands. Nitrogen mineralization during anaerobic incubations were about 10% greater on Albizia soils. Gross microbial mineralization and immobilization were determined by estimating the gross rates of N transformation by the ¹⁵N-isotope pool dilution techniques. Across species and sites, a strong linear positive relationship was obtained for gross immobilization and gross mineralization indicating faster gross immobilization as gross mineralization increases. Soil microbial biomass on Albizia soils contained larger proportion of it as bacterial biomass, while larger proportion of fungi biomass comprised the microbial biomass under Eucalyptus soils. This study clearly showed that the presence of Albizia increased total N pools and N supply to the ecosystem. The overall effect on soil fertility will need to be characterized by the effect of the N₂-fixer on other nutrients, especially the effect on phosphorus. Received: 28 February 1997 / Accepted: 22 September 1997     

The role of gross and net N transformation processes and NH4 + and NO3 − immobilization in controlling the mineral N pool of a temperate mixed deciduous forest soil

In many forests of Europe and north-eastern North America elevated N deposition has opened the forest N cycle, resulting in NO₃ ^? leaching. On the other hand, despite this elevated N deposition, the dominant fate of NO₃ ^? and NH₄ ⁺ in some of these forests is biotic or abiotic immobilization in the soil organic matter pool, preventing N losses. The environmental properties controlling mineral N immobilization and the variation and extent of mineral N immobilization in forest soils are not yet fully understood. In this study we investigated a temperate mixed deciduous forest, which is subjected to an average N deposition of 36.5 kg N ha^?1 yr^?1, but at the same time shows low NO₃ ^? concentrations in the groundwater. The aim of this study was to investigate whether the turnover rate of the mineral N pool could explain these low N leaching losses. A laboratory ¹⁵N pool dilution experiment was conducted to study gross and net N mineralization and nitrification and mineral N immobilization in the organic and uppermost (0–10 cm) mineral layer of the forest soil. Two locations, one at the forest edge (GE) and another one 145 m inside the forest (GF1), were selected. In the organic layers of GE and GF1, the gross N mineralization averaged 10.9 and 11.1 mg N kg^?1 d^?1, the net N mineralization averaged 6.1 and 6.8 mg N kg^?1 d^?1 and NH₄ ⁺ immobilization rates averaged 3.8 and 3.6 mg N kg^?1 d^?1. In the organic layer of GE and GF1, the average gross nitrification was 3.8 and 4.6 mg N kg^?1 d^?1, the average net nitrification was ?25.2 and ?31.3 mg N kg^?1 d^?1 and the NO₃ ^? immobilization rates averaged 29.0 and 35.9 mg N kg^?1 d^?1. For the mineral (0–10 cm) layer the same trend could be observed, but the N transformation rates were much lower for the NH₄ ⁺ pool and not significantly different from zero for the NO₃ ^? pool. Except for the turnover of the NH₄ ⁺ pool in the mineral layer, no significant differences were observed between location GE and GF1. The ratio of NH₄ ⁺ immobilization to gross N mineralization, gross N mineralization to gross nitrification, and NO₃ ^? immobilisation to gross nitrification led to the following observations. The NH₄ ⁺ pool of the forest soil was controlled by N mineralization and NO₃ ^? immobilization was importantly controlling the forest NO₃ ^? pool. Therefore it was concluded that this process is most probably responsible for the limited NO₃ ^? leaching from the forest ecosystem, despite the chronically high N deposition rates.     

Soil prokaryotic community shows no response to 2 years of simulated nitrogen deposition in an arid ecosystem in northwestern China

An arid ecosystem might be sensitive to nitrogen (N) deposition, but the associated ecosystem-specific response of soil microbes is not well studied. To assess the N enrichment effects on plant and prokaryotic community diversity, we performed a two-year NH₄NO₃ treatment in a desert steppe in northwestern China. Results showed that N addition increased plant aboveground biomass and decreased plant Shannon diversity. A C₄ herb (Salsola collina) became dominant, and loss of legume species was observed. The concentrations of soil NH₄⁺-N, NO₃⁻-N, microbial biomass N, and the plant aboveground biomass N pool increased in contrast to total N, suggesting that the N input into the arid ecosystem might mainly be assimilated by plants and exit the ecosystem. Remarkably, the α-diversity and structure of the soil prokaryotic community did not vary even at the highest N addition rate. Structural equation modelling further found that the plant aboveground N pool counteracted the acidification effect of N deposition and maintained soil pH thus partially stabilizing the composition of prokaryotic communities in a desert steppe. These findings suggested that the plants and N loss might contribute to the lack of responsiveness of soil prokaryotic community to N deposition in a desert steppe.     

Wildfire Effects on Soil Gross Nitrogen Transformation Rates in Coniferous Forests of Central Idaho, USA

Forest fires often result in a series of biogeochemical processes that increase soil nitrate (NO₃ ^?) concentrations for several years; however, the dynamic nature of inorganic nitrogen (N) cycling in the plant–microbe–soil complex makes it challenging to determine the direct causes of increased soil NO₃ ^?. We measured gross inorganic N transformation rates in mineral soils 2 years after wildfires in three central Idaho coniferous forests to determine the causes of the elevated soil NO₃ ^?. We also measured key factors that could affect the soil N processes, including temperature during soil incubation in situ, soil water content, pH and carbon (C) availability. We found no significant differences (P = 0.461) in gross nitrification rates between burned and control soils. However, microbial NO₃ ^? uptake rates were significantly lower (P = 0.078) in burned than control soils. The reduced consumption of NO₃ ^? caused slightly elevated NO₃ ^? concentrations in the burned soils. C availability was positively correlated with microbial NO₃ ^? uptake rates. Despite reduced microbial NO₃ ^? uptake capacity in the burned soils, soil microbes were a strong enough N sink to maintain low soil NO₃ ^? concentrations 2 years post fire. Soil NH₄ ⁺ concentrations between the treatments were not significantly different (P = 0.673). However, gross NH₄ ⁺ production and microbial uptake rates in burned soils were significantly lower (P = 0.028 and 0.035, respectively) than in the controls, and these rates were positively correlated with C availability. Our results imply that C availability is an important factor regulating soil N cycling of coniferous forests in the region.     

Variability of soil N cycling and N2O emission in a mixed deciduous forest with different abundance of beech

The mixture of other broadleaf species into beech forests in Central Europe leads to an increase of tree species diversity, which may alter soil biochemical processes. This study was aimed at 1) assessing differences in gross rates of soil N cycling among deciduous stands of different beech (Fagus sylvatica L.) abundance in a limestone area, 2) analyzing the relationships between gross rates of soil N cycling and forest stand N cycling, and 3) quantifying N₂O emission and determining its relationship with gross rates of soil N cycling. We used ¹⁵N pool dilution techniques for soil N transformation measurement and chamber method for N₂O flux measurement. Gross rates of mineral N production in the 0–5 cm mineral soil increased across stands of decreasing beech abundance and increasing soil clay content. These rates were correlated with microbial biomass which, in turn, was influenced by substrate quantity, quality and soil fertility. Leaf litter-N, C:N ratio and base saturation in the mineral soil increased with decreasing beech abundance. Soil mineral N production and assimilation by microbes were tightly coupled, resulting in low N₂O emissions. Annual N₂O emissions were largely contributed by the freeze-thaw event emissions, which were correlated with the amount of soil microbial biomass. Our results suggest that soil N availability may increase through the mixture of broadleaf species into beech forests.     

Microbial biomass and nitrogen cycling responses to fertilization and litter removal in young northern hardwood forests

The influence of site fertility on soil microbial biomass and activity is not well understood but is likely to be complex because of interactions with plant responses to nutrient availability. We examined the effects of long-term (8 yr) fertilization and litter removal on forest floor microbial biomass and N and C transformations to test the hypothesis that higher soil resource availability stimulates microbial activity. Microbial biomass and respiration decreased by 20–30 % in response to fertilization. Microbial C averaged 3.8 mg C/g soil in fertilized, 5.8 mg C/g in control, and 5.5 mg C/g in litter removal plots. Microbial respiration was 200 µg CO₂-C g^–1 d^–1 in fertilized plots, compared to 270 µg CO₂-C g^–1 d^–1 in controls. Gross N mineralization and N immobilization did not differ among treatments, despite higher litter nutrient concentrations in fertilized plots and the removal of substantial quantities of C and N in litter removal plots. Net N mineralization was significantly reduced by fertilization. Gross nitrification and NO₃ ^– immobilization both were increased by fertilization. Nitrate thus became a more important part of microbial N cycling in fertilized plots even though NH₄ ⁺ availability was not stimulated by fertilization.Soil microorganisms did not mineralize more C or N in response to fertilization and higher litter quality; instead, results suggest a difference in the physiological status of microbial biomass in fertilized plots that influenced N transformations. Respiration quotients (qCO₂, respiration per unit biomass) were higher in fertilized plots (56 µg CO₂-C mg C^–1 d^–1) than control (48 µg CO₂-C mg C^–1 d ^–1) or litter removal (45 µg CO₂-C mg C^–1 d^–1), corresponding to higher microbial growth efficiency, higher proportions of gross mineralization immobilized, and lower net N mineralization in fertilized plots. While microbial biomass is an important labile nutrient pool, patterns of microbial growth and turnover were distinct from this pool and were more important to microbial function in nitrogen cycling.     

Controls over Soil Nitrogen Pools in a Semiarid Grassland Under Elevated CO2 and Warming

Long-term responses of terrestrial ecosystems to the combined effects of warming and elevated CO₂ (eCO₂) will likely be regulated by N availability. The stock of soil N determines availability for organisms, but also influences loss to the atmosphere or groundwater. eCO₂ and warming can elicit changes in soil N via direct effects on microbial and plant activity, or indirectly, via soil moisture. Detangling the interplay of direct- and moisture-mediated impacts on soil N and the role of organisms in controlling soil N will improve predictions of ecosystem-level responses. We followed individual soil N pools over two growing seasons in a semiarid temperate grassland, at the Prairie Heating and CO₂ Enrichment experiment. We evaluated relationships of N pools with environmental factors and explored the role of plants by assessing plant biomass, plant N, and plant inputs to soil. We also assessed N forms in plots with and without vegetation to remove plant-mediated effects. Our study demonstrated that the effects of warming and eCO₂ are highly dependent on individual N form and on year. In this water-constrained grassland, eCO₂, warming and their combination appear to impact soil N pools through a complex combination of direct- and moisture-mediated effects. eCO₂ decreased NO₃ ^? but had neutral to positive effects on NH₄ ⁺ and dissolved organic N (DON), particularly in a wet year. Warming increased NO₃ ^? availability due to a combination of indirect drying and direct temperature-driven effects. Warming also increased DON only in vegetated plots, suggesting plant mediation. Our results suggest that impacts of combined eCO₂ and warming are not always equivalent for plant and soil pools; although warming can help offset the decrease in NO₃ ^? availability for plants under eCO₂, the NO₃ ^? pool in soil is mainly driven by the negative effects of eCO₂.     

Effects of Bromus tectorum invasion on microbial carbon and nitrogen cycling in two adjacent undisturbed arid grassland communities

Soil nitrogen (N) is an important component in maintaining ecosystem stability, and the introduction of non-native plants can alter N cycling by changing litter quality and quantity, nutrient uptake patterns, and soil food webs. Our goal was to determine the effects of Bromus tectorum (C₃) invasion on soil microbial N cycling in adjacent non-invaded and invaded C₃ and C₄ native arid grasslands. We monitored resin-extractable N, plant and soil δ¹³C and δ¹⁵N, gross rates of inorganic N mineralization and consumption, and the quantity and isotopic composition of microbial phospholipid biomarkers. In invaded C₃ communities, labile soil organic N and gross and net rates of soil N transformations increased, indicating an increase in overall microbial N cycling. In invaded C₄ communities labile soil N stayed constant, but gross N flux rates increased. The δ¹³C of phospholipid biomarkers in invaded C₄ communities showed that some portion of the soil bacterial population preferentially decomposed invader C₃-derived litter over that from the native C₄ species. Invasion in C₄ grasslands also significantly decreased the proportion of fungal to bacterial phospholipid biomarkers. Different processes are occurring in response to B. tectorum invasion in each of these two native grasslands that: 1) alter the size of soil N pools, and/or 2) the activity of the microbial community. Both processes provide mechanisms for altering long-term N dynamics in these ecosystems and highlight how multiple mechanisms can lead to similar effects on ecosystem function, which may be important for the construction of future biogeochemical process models.     

Effects of nitrogen addition on vegetation and ecosystem carbon in a semi-arid grassland

To clarify responses of plant and soil carbon (C) and nitrogen (N) pools in grassland ecosystem to N addition, a field experiment was performed in a grassland in Keerqin Sandy Lands, Northeast China. We investigated vegetation composition and C and N pools of plant and soil (0–30 cm) after five consecutive years of N addition at a rate of 20 g N m^?2 y^?1. Vegetation composition and species diversity responded dramatically to N addition, as dominance by C₄ perennials was replaced with C₃ annuals. Carbon in aboveground pool increased significantly (over two-fold), mainly due to the increase of the C in aboveground living plants and surface litter, which increased by 98 and 134%, respectively. Although soil C did not change significantly, the root C pool decreased in response to 5 years of N addition. The total ecosystem C pool was not significantly impacted by N addition because the large soil pool did not respond to N addition, and the increase in aboveground C was offset by the decrease in root C pool. Moreover, N addition significantly increased the aboveground N pool, but had no significant effects on belowground and total ecosystem N pools. Our results suggest that in the mid-term N addition alters the C and N partitioning in above- and belowground pools, but has no significant effects on total ecosystem C and N pools in these N-limited grasslands.     

Long-term movement of 15N tracers into fine woody debris under chronically elevated N inputs

Two key questions in the study of large-scale C (carbon) and N (nitrogen) cycling in temperate forests are how N cycling in soil detritus controls ecosystem-level retention of elevated N deposition, and whether elevated N deposition is likely to cause increases in C pools. The large C:N ratios in woody detritus make it a potentially important contributor to N retention, if N immobilization increases, and a potentially important contributor to C sequestration, if pool sizes increase. We studied N concentrations, C:N ratios, and pool sizes of N and biomass in fine woody debris (FWD < 5 cm diam.) 12 years into a long-term N-amendment study in two contrasting forests, a naturally-regenerated forest dominated by Quercus spp., and a 63-yr old plantation of Pinus resinosa. We also quantitatively recovered ¹⁵N tracers (originally applied as ¹⁵NH₄ and ¹⁵NO₃) in FWD, eight years following their application in the same study, in both ambient and N-amended plots. We used these data to test predictions of tracer redistributions made by a biogeochemical process model that included ¹⁵N. Results from the N pool-size analysis and the ¹⁵N tracer-recovery analysis indicated that under elevated N inputs of 5 g N m^–2 yr^–1 (as NH₄NO₃) over the decadal time period, only 0.15%–0.76% of the elevated N inputs were recovered in FWD of N-amended plots relative to ambient. Any increase in N immobilization in wood appeared to be minimal, in agreement with model predictions. Under N amendments, pool sizes of C in FWD were not significantly different from ambient, whereas pool sizes of N were marginally higher. Patterns of ¹⁵NH₄ vs. ¹⁵NO₃ recovery, treatment differences, and forest-type differences suggested that plant uptake, rather than detrital immobilization, was the dominant mechanism of ¹⁵N tracer movement into FWD. This result indicates that plant-soil cycling operating over a decadal time scale or longer controls C:N ratios and N pool sizes in woody debris.     

Effects of long-term experimental warming on plants and soil microbes in a cool temperate semi-natural grassland in Japan

To clarify the effects of long-term warming on ecosystem matter cycling, we conducted an in situ 7-year experimental warming (2009–2015) using infrared heaters in a cool temperate semi-natural grassland in Japan. We measured plant aboveground biomass, soil total C and N, soil inorganic N (NH₄ ⁺-N and NO₃ ^?-N), and soil microbial biomass for 7 years (2009–2015). We also measured heterotrophic respiration for 2 years (2013–2014) and assessed net N mineralization and nitrification in 2015. We found that warming immediately increased plant aboveground biomass, but this effect ceased in 2013. However, the soil microbial biomass was continuously depressed by warming. Soil inorganic N concentrations in warmed plots substantially increased in the later years of the experiment (2013–2015) and the potential net N mineralization rate was also higher than in the earlier years. In contrast, heterotrophic respiration decreased with warming in 2013–2014. Our observations indicate that long-term warming has a contrasting effect on plants and soil microbes. In addition, the warming could have different effects on subterranean C and N cycling. To enhance the accuracy of estimation of future climate change, it is essential to continuously observe the warming effects on ecosystems and to focus on the change in subterranean C and N cycling.     

Interactions between plant growth and soil nutrient cycling under elevated CO2: a meta-analysis

free air carbon dioxide enrichment (FACE) and open top chamber (OTC) studies are valuable tools for evaluating the impact of elevated atmospheric CO₂ on nutrient cycling in terrestrial ecosystems. Using meta‐analytic techniques, we summarized the results of 117 studies on plant biomass production, soil organic matter dynamics and biological N₂ fixation in FACE and OTC experiments. The objective of the analysis was to determine whether elevated CO₂ alters nutrient cycling between plants and soil and if so, what the implications are for soil carbon (C) sequestration. Elevated CO₂ stimulated gross N immobilization by 22%, whereas gross and net N mineralization rates remained unaffected. In addition, the soil C : N ratio and microbial N contents increased under elevated CO₂ by 3.8% and 5.8%, respectively. Microbial C contents and soil respiration increased by 7.1% and 17.7%, respectively. Despite the stimulation of microbial activity, soil C input still caused soil C contents to increase by 1.2% yr^?1. Namely, elevated CO₂ stimulated overall above‐ and belowground plant biomass by 21.5% and 28.3%, respectively, thereby outweighing the increase in CO₂ respiration. In addition, when comparing experiments under both low and high N availability, soil C contents (+2.2% yr^?1) and above‐ and belowground plant growth (+20.1% and+33.7%) only increased under elevated CO₂ in experiments receiving the high N treatments. Under low N availability, above‐ and belowground plant growth increased by only 8.8% and 14.6%, and soil C contents did not increase. Nitrogen fixation was stimulated by elevated CO₂ only when additional nutrients were supplied. These results suggest that the main driver of soil C sequestration is soil C input through plant growth, which is strongly controlled by nutrient availability. In unfertilized ecosystems, microbial N immobilization enhances acclimation of plant growth to elevated CO₂ in the long‐term. Therefore, increased soil C input and soil C sequestration under elevated CO₂ can only be sustained in the long‐term when additional nutrients are supplied.