Global pattern of organic carbon pools in forest soils.

Understanding the mechanisms of soil organic carbon (SOC) sequestration in forests is vital to ecosystem carbon budgeting and helps gain insight in the functioning and sustainable management of world forests. An explicit knowledge of the mechanisms driving global SOC sequestration in forests is still lacking because of the complex interplays between climate, soil, and forest type in influencing SOC pool size and stability. Based on a synthesis of 1179 observations from 292 studies across global forests, we quantified the relative importance of climate, soil property, and forest type on total SOC content and the specific contents of physical (particulate vs. mineral-associated SOC) and chemical (labile vs. recalcitrant SOC) pools in upper 10 cm mineral soils, as well as SOC stock in the O horizons. The variability in the total SOC content of the mineral soils was better explained by climate (47%-60%) and soil factors (26%-50%) than by NPP (10%-20%). The total SOC content and contents of particulate (POC) and recalcitrant SOC (ROC) of the mineral soils all decreased with increasing mean annual temperature because SOC decomposition overrides the C replenishment under warmer climate. The content of mineral-associated organic carbon (MAOC) was influenced by temperature, which directly affected microbial activity. Additionally, the presence of clay and iron oxides physically protected SOC by forming MAOC. The SOC stock in the O horizons was larger in the temperate zone and Mediterranean regions than in the boreal and sub/tropical zones. Mixed forests had 64% larger SOC pools than either broadleaf or coniferous forests, because of (i) higher productivity and (ii) litter input from different tree species resulting in diversification of molecular composition of SOC and microbial community. While climate, soil, and forest type jointly determine the formation and stability of SOC, climate predominantly controls the global patterns of SOC pools in forest ecosystems.

Differential responses of soil phosphorus fractions to varied nitrogen compound additions in a meadow steppe.

Nitrogen (N) addition can greatly influence soil inorganic phosphorus (Pi) and organic phosphorus (Po) transformations. However, whether and how the N compound forms may differentially affect the soil P fractions remain unclear. Here, we investigated the responses of soil Pi (labile Pi, moderately-occluded Pi, and recalcitrant Pi) and Po fractions (labile Po and stable Po) to varying addition rates of three N compounds ((NH4)2SO4, NH4NO3, and urea) in a meadow steppe in northern China. Our studies revealed that with increasing N addition rate, soil labile and moderately-occluded Pi increased, accompanied by decreases in soil recalcitrant Pi. This shift was attributed to N-induced soil acidification, which accelerated the conversion of recalcitrant Pi into labile and moderately-occluded Pi. Soil labile Po decreased with increasing rate of N addition, whilst soil stable Po was not affected. Regardless of the compound forms, N addition increased soil Olsen-P, suggesting a potential alleviation of P limitation in this grassland ecosystem. The effect of N addition on soil labile Pi was significantly greater with addition of urea than with addition of either (NH4)2SO4 or NH4NO3, indicating that urea was more efficient in enhancing soil P availability. Addition of (NH4)2SO4 imposed a more pronounced positive effect on soil moderately-occluded Pi than the addition of either NH4NO3 or urea, mainly due to the greater mobilization of recalcitrant Pi as a result of higher soil acidification strength of (NH4)2SO4. These findings underscore the importance of considering the distinct effects of different N compounds when studying grassland soil P dynamics and availability in response to N addition.

Enhanced foliar 15 N enrichment with increasing nitrogen addition rates: Role of plant species and nitrogen compounds.

Determining the abundance of N isotope (δ15 N) in natural environments is a simple but powerful method for providing integrated information on the N cycling dynamics and status in an ecosystem under exogenous N inputs. However, whether the input of different N compounds could differently impact plant growth and their 15 N signatures remains unclear. Here, the response of 15 N signatures and growth of three dominant plants (Leymus chinensis, Carex duriuscula, and Thermopsis lanceolata) to the addition of three N compounds (NH4 HCO3 , urea, and NH4 NO3 ) at multiple N addition rates were assessed in a meadow steppe in Inner Mongolia. The three plants showed different initial foliar δ15 N values because of differences in their N acquisition strategies. Particularly, T. lanceolata (N2 -fixing species) showed significantly lower 15 N signatures than L. chinensis (associated with arbuscular mycorrhizal fungi [AMF]) and C. duriuscula (associated with AMF). Moreover, the foliar δ15 N of all three species increased with increasing N addition rates, with a sharp increase above an N addition rate of ~10 g N m-2  year-1 . Foliar δ15 N values were significantly higher when NH4 HCO3 and urea were added than when NH4 NO3 was added, suggesting that adding weakly acidifying N compounds could result in a more open N cycle. Overall, our results imply that assessing the N transformation processes in the context of increasing global N deposition necessitates the consideration of N deposition rates, forms of the deposited N compounds, and N utilization strategies of the co-existing plant species in the ecosystem.

Nitrogen addition and mowing had only weak interactive effects on macronutrients in plant-soil systems of a typical steppe in Inner Mongolia.

Effective management of macronutrients is pivotal in the optimization and provisioning of ecosystem services in grassland areas, particularly in degraded grasslands. In such instances where mowing and nitrogen (N) fertilization have emerged as predominant management strategies, nutrient management is especially important. However, the precise effects of these concurrent practices on the distribution of macronutrients in plant-soil systems remain unclear. Here we evaluated the effects of 12 years of N addition (2, 10, and 50 g N m-2 year-1) and mowing on the concentrations and pools of six macronutrients (i.e., N; phosphorus P; sulfur S, calcium Ca, magnesium Mg, and potassium K) in three plant components (aboveground plants, litter, and belowground roots) at the community level and in the soil in a typical steppe in Inner Mongolia. Our results revealed that N addition generally raised the N concentration in the entire plant-soil system, regardless of whether plots were mowed. Higher N addition (10 and 50 g N m-2 year-1) also led to higher concentrations of P (+22%, averaging two N addition rates), S (+16%), K (+22%), Ca (+22%), and Mg (+24%) in plants but lower concentrations of these nutrients in the litter. Similar decreases in K (-9%), Ca (-46%), and Mg (-8%) were observed in the roots. In light of the observed increases in vegetation biomass and the lack of pronounced changes in soil bulk density, we found that the ecosystem N enrichment resulted in increased pools of all measured macronutrients in plants, litter, and roots (with the exception of Ca in the roots) while concurrently decreased the pools of P (-20%, averaging two higher N addition rates), S (-12%), K (-10%), Ca (-37%), and Mg (-19%) in the soil, with no obvious effect of the mowing practice. Overall, mowing exhibited a very limited capacity to alleviate the effects of long-term N addition on macronutrients in the plant-soil system. These findings highlight the importance of considering the distribution of macronutrients across distinct plant organs and the dynamic nutrient interplay between plants and soil, particularly in the context of long-term fertilization and mowing practices, when formulating effective grassland management strategies.

A novel 13 C pulse-labelling method to quantify the contribution of rhizodeposits to soil respiration in a grassland exposed to drought and nitrogen addition.

Rhizodeposition plays an important role in below-ground carbon (C) cycling. However, quantification of rhizodeposition in intact plant-soil systems has remained elusive due to methodological issues. We used a 13 C-CO2 pulse-labelling method to quantify the contribution of rhizodeposition to below-ground respiration. Intact plant-soil cores were taken from a grassland field, and in half, shoots and roots were removed (unplanted cores). Both unplanted and planted cores were assigned to drought and nitrogen (N) treatments. Afterwards, shoots in planted cores were pulse labelled with 13 C-CO2 and then clipped to determine total below-ground respiration and its δ13 C. Simultaneously, δ13 C was measured for the respiration of live roots, soils with rhizodeposits, and unplanted treatments, and used as endmembers with which to determine root respiration and rhizodeposit C decomposition using two-source mixing models. Rhizodeposit decomposition accounted for 7-31% of total below-ground respiration. Drought reduced decomposition of both rhizodeposits and soil organic carbon (SOC), while N addition increased root respiration but not the contribution of rhizodeposit C decomposition to below-ground respiration. This study provides a new approach for the partitioning of below-ground respiration into different sources, and indicates that decomposition of rhizodeposit C is an important component of below-ground respiration that is sensitive to drought and N addition in grassland ecosystems.

Increasing rates of long-term nitrogen deposition consistently increased litter decomposition in a semi-arid grassland.

The continuing nitrogen (N) deposition observed worldwide alters ecosystem nutrient cycling and ecosystem functioning. Litter decomposition is a key process contributing to these changes, but the numerous mechanisms for altered decomposition remain poorly identified. We assessed these different mechanisms with a decomposition experiment using litter from four abundant species (Achnatherum sibiricum, Agropyron cristatum, Leymus chinensis and Stipa grandis) and litter mixtures representing treatment-specific community composition in a semi-arid grassland under long-term simulation of six different rates of N deposition. Decomposition increased consistently with increasing rates of N addition in all litter types. Higher soil manganese (Mn) availability, which apparently was a consequence of N addition-induced lower soil pH, was the most important factor for faster decomposition. Soil C : N ratios were lower with N addition that subsequently led to markedly higher bacterial to fungal ratios, which also stimulated litter decomposition. Several factors contributed jointly to higher rates of litter decomposition in response to N deposition. Shifts in plant species composition and litter quality played a minor role compared to N-driven reductions in soil pH and C : N, which increased soil Mn availability and altered microbial community structure. The soil-driven effect on decomposition reported here may have long-lasting impacts on nutrient cycling, soil organic matter dynamics and ecosystem functioning.

Intensity and frequency of nitrogen addition alter soil chemical properties depending on mowing management in a temperate steppe.

Anthropogenic nitrogen (N) enrichment can significantly alter soil chemical properties in various ecosystems. Previous manipulative N experiments mainly focused on the intensity of N addition on soil properties by changing N input rates. It remains unclear, however, whether frequency of N addition can affect soil chemical properties. We examined the effects of frequency (2 versus 12 applications yr-1) and rate (ranging from 0 to 50 g N m-2 yr-1) of N addition on soil chemical properties of pH, base cations, soil pH buffering capacity (pHBC), and soil available micronutrients in a temperate steppe with and without mowing. Mowing significantly increased the effective cation exchange capacity (ECEC), soil exchangeable Ca and Na, available Fe, and soil pHBC when N was applied at low frequency. Low frequency of N addition significantly decreased soil pH and exchangeable Na but increased soil exchangeable Mg without mowing; however, it increased soil exchangeable Na and available Zn with mowing, while available Fe and Mn increased both with and without mowing. Higher rates of N addition (≥20 g N m-2 yr-1) decreased soil pH, ECEC and exchangeable Ca but increased soil available Fe, Mn and Cu regardless of the mowing treatment and frequency of N addition. Changes in soil organic matter, pHBC and ECEC were the main reasons affecting soil pH across mowing and N application treatments. Our results indicate that frequency of N addition played an essential role in altering soil chemical properties. Simulating N deposition via large and infrequent N additions can underestimate (exchangeable Mg and available Fe and Mn) or overestimate (soil pH and exchangeable Na) changes in soil properties. Our results further suggest that the effects of frequency of N addition on soil chemical attributes in semi-arid grassland ecosystems can be regulated by appropriate mowing management.

Effects of nitrogen and water addition on trace element stoichiometry in five grassland species.

A 9-year manipulative experiment with nitrogen (N) and water addition, simulating increasing N deposition and changing precipitation regime, was conducted to investigate the bioavailability of trace elements, iron (Fe), manganese (Mn), copper (Cu), and zinc (Zn) in soil, and their uptake by plants under the two environmental change factors in a semi-arid grassland of Inner Mongolia. We measured concentrations of trace elements in soil and in foliage of five common herbaceous species including 3 forbs and 2 grasses. In addition, bioaccumulation factors (BAF, the ratio of the chemical concentration in the organism and the chemical concentration in the growth substrate) and foliar Fe:Mn ratio in each plant was calculated. Our results showed that soil available Fe, Mn and Cu concentrations increased under N addition and were negatively correlated with both soil pH and cation exchange capacity. Water addition partly counteracted the positive effects of N addition on available trace element concentrations in the soil. Foliar Mn, Cu and Zn concentrations increased but Fe concentration decreased with N addition, resulting in foliar elemental imbalances among Fe and other selected trace elements. Water addition alleviated the effect of N addition. Forbs are more likely to suffer from Mn toxicity and Fe deficiency than grass species, indicating more sensitivity to changing elemental bioavailability in soil. Our results suggested that soil acidification due to N deposition may accelerate trace element cycling and lead to elemental imbalance in soil-plant systems of semi-arid grasslands and these impacts of N deposition on semi-arid grasslands were affected by water addition. These findings indicate an important role for soil trace elements in maintaining ecosystem functions associated with atmospheric N deposition and changing precipitation regimes in the future.

Responses of Soil Bacterial Communities to Nitrogen Deposition and Precipitation Increment Are Closely Linked with Aboveground Community Variation.

It has been predicted that precipitation and atmospheric nitrogen (N) deposition will increase in northern China; yet, ecosystem responses to the interactive effects of water and N remain largely unknown. In particular, responses of belowground microbial community to projected global change and their potential linkages to aboveground macro-organisms are rarely studied. In this study, we examined the responses of soil bacterial diversity and community composition to increased precipitation and multi-level N deposition in a temperate steppe in Inner Mongolia, China, and explored the diversity linkages between aboveground and belowground communities. It was observed that N addition caused the significant decrease in bacterial alpha-diversity and dramatic changes in community composition. In addition, we documented strong correlations of alpha- and beta-diversity between plant and bacterial communities in response to N addition. It was found that N enriched the so-called copiotrophic bacteria, but reduced the oligotrophic groups, primarily by increasing the soil inorganic N content and carbon availability and decreasing soil pH. We still highlighted that increased precipitation tended to alleviate the effects of N on bacterial diversity and dampen the plant-microbe connections induced by N. The counteractive effects of N addition and increased precipitation imply that even though the ecosystem diversity and function are predicted to be negatively affected by N deposition in the coming decades; the combination with increased precipitation may partially offset this detrimental effect.

Weathering of pyrogenic organic matter induces fungal oxidative enzyme response in single culture inoculation experiments.

The addition of pyrogenic organic matter (PyOM), the aromatic carbon-rich product of the incomplete combustion of plant biomass or fossil fuels, to soil can influence the rate of microbial metabolism of native soil carbon. The interaction of soil heterotrophs with PyOM may be governed by the surficial chemical and physical properties of PyOM that evolve with environmental exposure. We present results of a 36-day laboratory incubation investigating the interaction of a common white-rot fungus, Trametes versicolor, with three forms of 13C-enriched (2.08 atom% 13C) PyOM derived from Pinus ponderosa (450 °C): one freshly produced, and two artificially weathered (254 nm, UV light-water treatment and water-leaching alone). Analysis (FTIR, XPS) of the UV-weathered PyOM showed increased aliphatic C-H content and oxidation of aromatic carbon relative to both the original and water-leached PyOM. The addition of both weathered forms of PyOM stimulated (positively primed) fungal respiration of the growth media, while the unaltered PyOM mildly inhibited (negatively primed) respiration. Artificial weathering resulted in higher oxidative (laccase and peroxidase) enzyme activity than unaltered PyOM, possibly the result of a diminished capacity to bind reactive substrates and extracellular enzymes after weathering. However, and contrary to expectations, simple water-leached weathering resulted in a relatively higher enzyme activity and respiration than that of UV-weathering. The 13C content of respired CO2 indicated negligible fungal oxidation of PyOM for all treatments, demonstrating the overall low microbial reactivity of this high temperature PyOM. The increased enzymatic and positive priming response of T. versicolor to weathered PyOM highlights the importance of weathering-induced chemistry in controlling PyOM-microbe-soil carbon interactions.