high-N Addition Research Articles

Determinants of tree population temporal stability in a temperate mixed forest over a gradient of nitrogen addition

Nitrogen (N) deposition is a significant threat to the functioning of forests and negatively impacts the delivery of forest goods and services. Contemporary management approaches seek to adapt forests to such N-deposition stressors, but to date how plant populations in natural forests respond to N deposition and what factors determine the contrasting responses among populations are still unclear. Here, we investigated the impact of N-addition (control: 0 kg ha−1 yr−1; low: 25 kg ha−1 yr−1; medium: 50 kg ha−1 yr−1; high: 75 kg ha−1 ha yr−1) on tree population temporal stability and how initial tree size, mycorrhizal type, and leaf N content (LNC; as a surrogate for functional trait composition) mediate tree population responses to N-addition in a Korean pine and mixed broadleaved dominated temperate forest in northern China. We quantified tree species population temporal stability as the ratio of mean to standard deviation of the year-by-year stem increments recorded in individual trees from 2015 to 2022 experimental period. The results showed different temporal stabilities of tree species among four N-addition levels, with the highest population stability observed within the high N-addition plots. Furthermore, initial tree size had significantly (p < 0.001) positive effects on population temporal stability. The effect of LNC and initial tree size were also contingent on the level of N applied. Specifically, increase in tree population LNC reduced population temporal stability in all plots where N was added. Our results imply that retention of large-sized trees and species with resource-conservative strategies (e.g., low LNC) could enhance forest stability under N deposition.

Canopy and understory nitrogen additions differently affect soil microbial residual carbon in a temperate forest.

Atmospheric nitrogen (N) deposition in forests can affect soil microbial growth and turnover directly through increasing N availability and indirectly through altering plant-derived carbon (C) availability for microbes. This impacts microbial residues (i.e., amino sugars), a major component of soil organic carbon (SOC). Previous studies in forests have so far focused on the impact of understory N addition on microbes and microbial residues, but the effect of N deposition through plant canopy, the major pathway of N deposition in nature, has not been explicitly explored. In this study, we investigated whether and how the quantities (25 and 50 kg N ha-1 year-1) and modes (canopy and understory) of N addition affect soil microbial residues in a temperate broadleaf forest under 10-year N additions. Our results showed that N addition enhanced the concentrations of soil amino sugars and microbial residual C (MRC) but not their relative contributions to SOC, and this effect on amino sugars and MRC was closely related to the quantities and modes of N addition. In the topsoil, high-N addition significantly increased the concentrations of amino sugars and MRC, regardless of the N addition mode. In the subsoil, only canopy N addition positively affected amino sugars and MRC, implying that the indirect pathway via plants plays a more important role. Neither canopy nor understory N addition significantly affected soil microbial biomass (as represented by phospholipid fatty acids), community composition and activity, suggesting that enhanced microbial residues under N deposition likely stem from increased microbial turnover. These findings indicate that understory N addition may underestimate the impact of N deposition on microbial residues and SOC, highlighting that the processes of canopy N uptake and plant-derived C availability to microbes should be taken into consideration when predicting the impact of N deposition on the C sequestration in temperate forests.

Distinct roles of bacteria and fungi in mediating soil extracellular enzymes under long-term nitrogen deposition in temperate plantations

Soil microbes decompose soil organic matter by producing various extracellular enzymes, and also regulate soil carbon (C) and nutrient cycles. However, how shifts in soil microbial community under long-term nitrogen (N) deposition govern soil extracellular enzyme activities (EEAs) is not known. Here, we conducted a decade-long N fertilization experiment consisting of control, low-N (2 g N m−2 year−1), medium-N (5 g N m−2 year−1), and high-N (10 g N m−2 year−1) treatments in a N-limited temperate plantation in northern China. The diversity and community composition of fungi and bacteria were determined with bacterial 16S rRNA genes and fungal internal transcribed spacer genes sequencing. Soil EEAs involved in C, N, and phosphorus (P) cycles were also measured. The results showed that high-N addition increased fungal diversity and altered microbial community composition. In particular, rare fungal diversity and rare bacterial community composition were more sensitive to N addition. N addition improved β-1,4-glucosidase (BG) and acid phosphatase (AP) activities by altering bacterial community composition, and inhibited leucine aminopeptidase (LAP) and polyphenolic oxidase (PPO) activities by affecting fungal functional groups. Specifically, shifts in bacterial community composition caused by N addition were reflected in the significant increase in the dominant class Thermoleophilia, which could be an indicator of secreted hydrolase involved in C and P cycles. Symbiotrophic fungi were more closely associated with LAP for acquiring N and PPO. Our data provide a novel view of the link between major extracellular enzymes and specific microbial taxa under the background of N deposition.

Effects of 12-Year Nitrogen Addition and Mowing on Plant-Soil Micronutrients in a Typical Steppe.

Changes in soil micronutrient availability may have adverse consequences on grassland productivity, yet it’s still largely unclear how concurrent human practices, such as fertilization and mowing, affect micronutrient cycling in the plant-soil systems. Here, we measured six essential micronutrient (Fe, Mn, Cu, Zn, Co and Mo) contents in both plant pool (separated as aboveground plant parts, litter, and belowground roots) at the community level and soil pool (0−10 cm depth) after 12-year consecutive nitrogen (N) addition (0, 2, 10, and 50 g N m−2 year−1) and mowing in a typical steppe of the Mongolian Plateau. The results show that (i) medium-N (10 g m−2 year−1) and high-N (50 g m−2 year−1) addition rates significantly increased contents of soil-available Fe (+310.0%, averaging across the two N addition rates), Mn (+149.2%), Co (+123.6%) and Mo (+73.9%) irrespective of mowing treatment, whereas these addition treatments usually decreased contents of soil total Fe (−8.9%), Mn (−21.6%), Cu (−15.9%), Zn (−19.5%), Co (−16.4%) and Mo (−34.7%). (ii) Contents of Fe in aboveground plant parts, litter, and roots significantly decreased, whereas plant Mn increased with N addition. Contents of above ground plant Cu, Zn, Co, and Mo significantly decreased at high-N addition rate, whereas contents of micronutrients in roots and litters, except for Fe, generally increased with N addition. Moreover, the total amount of micronutrients in the plant pool (contents × biomass) significantly increased at the medium-N addition rate but decreased at the high-N addition rate. All N addition rates significantly enlarged the pool of litter micronutrients, and roots could hold more micronutrients under N addition, especially combined with mowing treatment. Importantly, although mowing could regulate the effects of N addition on variables (i) and (ii), the effects were weaker overall than those of N addition. (iii) Changes in root micronutrients, except for Mn, could explain corresponding changes in plant micronutrients (R2: 0.19−0.56, all p < 0.01), and significant linear correlations were also observed between soil-available Fe and Fe in plant and roots. Aboveground plant Mn was significantly correlated with soil-available Mn, while Co and Mo in roots were also significantly correlated with soil-available Co and Mo. These results indicate that soil micronutrient supply capacity may decrease due to a decrease in total micronutrient contents after long-term N addition and mowing. They also suggest that different magnitude responses of soil micronutrients in plants (i.e., litters, roots) and soil should be considered when comprehensively examining nutrient cycling in grassland ecosystems.

Long-term nitrogen addition increased soil microbial carbon use efficiency in subalpine forests on the eastern edge of the Qinghai–Tibet Plateau

AimsNitrogen (N) deposition increased forest carbon (C) sink significantly, hence exploring the microscopic mechanisms is critical to predicting future global ecosystem C cycle, especially the effects of enhanced N deposition on soil microbial carbon use efficiency (CUE), which still unclear.MethodsWe evaluated the responses of soil microbial CUE to long-term (5 years) N addition in an evergreen broad-leaved forest and a mature coniferous forest by using a 13C isotope tracing method.ResultsThe results showed that the soil microbial CUE ranged from 0.38 to 0.51, which was smaller than the results obtained from the previous studies based the same method and forest type. In evergreen broad-leaved forest, the microbial CUE had no significant changes in the low N-addition treatment, but it was increased by 9.23% and 12.69% in medium and high N-addition treatments compared to the control. In coniferous forest, soil microbial CUE was increased by 14.64%, 21.89% and 24.34% in low, medium and high N-addition treatments, respectively. Moreover, the soil C:P and N:P are negatively relate to soil microbial CUE.ConclusionsOur findings indicate that the enhancing N deposition can increase soil microbial CUE and ultimately promote C sequestration, especially in coniferous forest. The imbalance of soil stoichiometry is the main impact factor of CUE under N addition. However, we speculate that the key to increase forest soil microbial CUE is to promote the decomposition rate of litter and thus increase the available C content.

High nitrogen addition after the application of sewage sludge compost decreased the bioavailability of heavy metals in soil

Nitrogen (N) fertilizer is highly significant in agricultural production, but long-term N addition causes changes in quality indicators, such as soil organic matter (SOM), thus affecting the absorption and accumulation of organic pollutants. Therefore, paying more attention to organic fertilizers in the development of green agriculture is necessary. However, the accumulation of heavy metals (HMs) contained in organic fertilizers (especially sewage sludge compost (SSC)) in the soil can cause environmental contamination, but how this cumulative reaction changes with the long-term N addition remains unclear. Here the SSC impact on the bioavailability of five typical HMs (cadmium–Cd, chromium–Cr, copper–Cu, lead–Pb and arsenic–As) in the soil-plant system before and after SSC application was demonstrated through a field study in soils with different application rates of 0, 100 and 300 kg N ha−1yr−1, respectively. Our results showed that SSC application increased the concentration of most HMs in soil profiles and plant systems (wheat roots and grains), but the accumulation rate of HMs and most bioaccumulation values (BAC–bioaccumulation coefficient and BCF–bioconcentration factor) in plant systems were both lower in high-N addition soil than that in the low-N group. Moreover, speciation distribution results further indicated that SSC application increased the LB (liable available form, including F1–water soluble, F2–ion exchangeable, and F3–bound to carbonates) form of HMs and decreased the PB (potentially available form, including F4–humic acids and F6–fraction bound to organic matter) form of HMs in high-N addition soil, respectively. Those results suggested that HM bioavailability in high-N addition soil was lower than that in low-N addition soil when applied with SSC. Overall, this study found that increasing soil N content can inhibit the bioavailability of HMs when applying SSC, providing suggestions for optimizing the trialability and risk assessment of SSC application.

Do long-term high nitrogen inputs change the composition of soil dissolved organic matter in a primary tropical forest?

Dissolved organic matter (DOM) plays a key role in forest carbon biogeochemistry by linking soil organic carbon (SOC) sequestration and water fluxes, which is further shaped by elevated atmospheric nitrogen (N) deposition. Although enhanced SOC sequestration was evidenced in tropical forests due to rising N deposition, it remains unclear how long-term N inputs affect soil DOM composition, which regulates SOC sequestration capability due to its mobility and biological instability. Here, the quantity, optical properties, and molecular-level characteristics of soil DOM based on a simulative N deposition experiment with four N addition levels (0, 5, 10, and 15 g m−2 yr−1) were studied in a primary tropical forest in south China. Results showed that 18 year N additions significantly altered soil DOM composition, with an increasing trend in soil dissolved organic carbon content. Medium- (10 g m−2 yr−1) and high-N addition (15 g m−2 yr−1) markedly elevated DOM average molecular weight by 12% and aromaticity, with specific ultraviolet absorbance at 254 nm increasing by 17%, modified aromatic index by 35%, and condensed aromatics by 67%. Medium- and high-N addition also increased recalcitrant DOM components but decreased other DOM components, with increasing percentages of lignin-like, tannin-like, and carboxylic-rich alicyclic molecule-like compounds, and decreasing percentage of more bioavailable contributions with H/C ratio >1.5. Importantly, significant correlations of the SOC content of the heavy fraction with optical properties and with recalcitrant DOM components were observed. These findings suggest that long-term N additions may alter soil DOM composition in a way to benefit soil OC storage in the primary tropical forests. It merits focusing on the mechanisms to association of soil DOM dynamics with SOC sequestration.

Effects of Nitrogen Addition on Soil Microbial Functional Diversity and Extracellular Enzyme Activities in Greenhouse Cucumber Cultivation

Greenhouses, commonly used for vegetable production, are experiencing large nitrogen (N) inputs in North China, which leads to soil acidification, increases soil N availability, and affects microbial community structure and composition. However, it remains unclear how N enrichment influences soil microbial functional activities in this region. In this study, we conducted a two-year pot experiment in a greenhouse to evaluate the effects of four different rates of N addition (0, 334, 668, and 1002 kg N ha−1 year−1) on cucumber soil properties, extracellular enzyme activities, and community level physiological profiles (CLPP). We found that high-N addition (1002 kg N ha−1) caused a massive accumulation of inorganic nitrogen and soil acidification, which was not beneficial to soil microbial activities. The color development (AWCD) values for the metabolism of microbial carbon sources and the activities of soil extracellular enzymes also showed a significant decrease in high N(N3) treatment. Additionally, the activity of leucine aminopeptidase (LAP) and polyphenol oxidase (PPO) of N3 decreased by 36% and 50% compared to the N0 and could be a good predictor for microbial functional diversity and microbial biomass carbon (MBC). Structural equation modeling (SEM) confirmed that the reduction of microbial functional diversity is mainly coregulated by the decline of soil pH and the change of cucumber BGB (belowground biomass) resulting from soil C and N imbalance. Overall, excessive N-fertilizer amendment can be more dangerous to microbial community functional diversity, especially for carbohydrate utilization which adversely affects cucumber yield in current intensive management.

Ecosystem function responses to nutrient enrichment mediated by mud content in soft sediment habitats

ABSTRACT Estuaries are threatened with increasing sedimentation and nutrients, however research is limited on the interaction of these stressors on ecosystem functions. A field experiment adding nitrogen (N) fertiliser at high (600 g N m−2) and medium (150 g N m−2) concentrations and control plots was established in two sandy and two muddy sites (mud content 0–15%) to investigate context dependency of nitrogen enrichment on nutrient cycling and primary production. There was a significant interaction of mud x treatment on ammonium (NH4 +) fluxes. High N-addition plots had 133x greater dark NH4 + efflux than controls in muddy sediments, compared to 17x greater dark NH4 + efflux in sandy sediments. Net primary production increased with mud content, while photosynthetic efficiency of the microphytobenthos (GPPchl-a ) decreased above 4% mud. Mud content also affected the bivalve, Austrovenus stutchburyi (>30 mm), macrofaunal abundance and functional groups, and their influence on nutrient cycling. Thus minor increases in mud can alter the ecosystem response to nutrient enrichment directly through changes in sediment properties and indirectly by affecting the bivalves and macrofauna that influence these processes. This study highlights how multiple stressors interact to predispose estuaries to tipping points.

Canopy and understory nitrogen addition have different effects on fine root dynamics in a temperate forest: implications for soil carbon storage.

Elucidating the effects of atmospheric nitrogen (N) deposition on fine root dynamics and the potential underlying mechanisms is required to understand the changes in belowground and aboveground carbon storage. However, research on these effects in forests has mostly involved direct understory addition of N and has ignored canopy interception and processing of N. Here, we conducted a field experiment comparing the effects of canopy addition of N (CAN) with those of understory addition of N (UAN) at three N-addition rates (0, 25 and 50 kg N ha-1 yr-1 ) on fine root dynamics in a temperate deciduous forest. Fine root production and biomass were significantly higher with CAN than with UAN. At the same N-addition rate, increases in fine root production with CAN were at least two-fold greater than with UAN. At the high N-addition rate and relative to the control, fine root biomass was significantly increased by CAN (by 23.5%) but was significantly decreased by UAN (by 12.2%). Our results indicate that traditional UAN may underestimate the responses of fine root dynamics to atmospheric N deposition in forest ecosystems. Canopy N processes should be considered for more realistic assessments of the effects of atmospheric N deposition in forests.

Soil microbial community responses to long-term nitrogen addition at different soil depths in a typical steppe

Global nitrogen (N) deposition has been influencing the structure of soil microbial communities, which play fundamental roles in modifying most biogeochemical processes. However, to date, our understanding of how long-term N deposition affects soil microbial communities, particularly those in subsurface soil, is largely incomplete. In this study, we examined soil microbial phospholipid fatty acids at different depths in the soil profile, including in topsoil (0–10 cm), midsoil (30–40 cm), and subsoil (70–100 cm), in a 10-year N addition experimental site in a semiarid steppe. We collected soil samples at four N addition treatment levels, including 0, 2, 10, and 50 g m−2 year−1, which represented control, low-N, medium-N, and high-N inputs, respectively. Our results showed that N addition remarkably shifted soil microbial communities by increasing the relative abundances of bacteria and Gram-positive (GP) bacteria, and decreasing Gram-negative (GN) bacterial relative abundance across the three soil layers by affecting soil pH and NO3−-N, particularly under the medium- and high-N addition rates. Moreover, the effects of N addition tended to increase with the N addition rate but diminished with soil depth. Soil pH, NO3−-N, and NH4+-N were the most important driving factors for changes in microbial community composition in topsoil, whereas soil total N (TN) and total carbon were important in midsoil, and TN, dissolved organic N, and soil moisture were important in subsoil. The relative abundances of bacteria and GN bacteria also decreased, whereas those of fungi and GP bacteria increased with soil depth regardless of N addition. Soil TN and pH were the most important factors in shaping the vertical distribution of soil microbial communities. Our results suggest that both N addition and soil depth may cause similar microbial community shifts, through which fungi and GP bacteria become dominant, but through different mechanisms. These results suggest that it is important to distinguish among the N-deposition effects on soil microbial communities at different soil depths when building soil system models.

Nitrogen Additions Retard Nutrient Release from Two Contrasting Foliar Litters in a Subtropical Forest, Southwest China

Litter decomposition plays a critical role in regulating biogeochemical cycles in terrestrial ecosystems and is profoundly impacted by increasing atmospheric nitrogen (N) deposition. Here, a N manipulation experiment was conducted to explore the effects of N additions (0 kg N ha−1 yr−1, 20 kg N ha−1 yr−1 and 40 kg N ha−1 yr−1) on decay rates and nutrients release of two contrasting species, the evergreen and nutrient-poor Michelia wilsonii and the deciduous and nutrient-rich Camptotheca acuminata, using a litterbag approach at the western edge of the Sichuan Basin of China. The decay rate and the mineralization of N and phosphorus (P) was faster in nutrient-rich C. acuminata litter than in nutrient-poor M. wilsonii litter, regardless of N regimes. N additions tended to decrease the decay constant (k value) in M. wilsonii litter, but had no effect on C. acuminata litter. N additions had no significant effects on carbon (C) release of both litter types. N additions showed negative effects on N and P release of M. wilsonii litter, particularly in the late decomposition stage. Moreover, for C. acuminata litter, N additions did not affect N release, but retarded P release in the late stage. N additions did not affect the C:N ratio in both litter types. However, N additions—especially high-N addition treatments—tended to reduce C:P and N:P ratios in both species. The effect of N addition on N and P remaining was stronger in M. wilsonii litter than in C. acuminata litter. The results of this study indicate that N additions retarded the nutrients release of two foliar litters. Thus, rising N deposition might favor the retention of N and P via litter decomposition in this specific area experiencing significant N deposition.

Nonlinear responses of total belowground carbon flux and its components to increased nitrogen availability in temperate forests.

With the increased interest in allocating more carbon (C) belowground for C sequestration, total belowground C flux (TBCF) and its dynamics have become an important research topic. However, it remains uncertain whether TBCF responds nonlinearly to increased nitrogen (N) availability and how its main components (root and ectomycorrhizal (ECM) fungi) contribute to TBCF. We established a four-yearN addition experiment with control, low-N, medium-N, and high-N fertilization treatments in a N-limited temperate forest in northern China. We measured TBCF and its three main components including root respiration, ECM fungal respiration, and root production. The involved edaphic and plant factors were also measured. TBCF showed a nonlinear response to the increasing amounts of N addition, accelerated by 10.37% in low-N addition and restrained by 10.29% in high-N addition. Contrasting patterns of the contributions of root respiration and ECM fungal respiration to TBCF implies different strategies of investment in roots and ECM fungi under the different N-availability statuses. The ratio of production and respiration in roots under N addition was nearly 1:2, which indicated that when soil N availability increases, roots prefer to lose C by overflow respiration rather than fix C in new biomass. The low-N addition increased TBCF by directly increasing root respiration and indirectly increasing coarse root biomass. The medium-N addition positively affected TBCF by increasing root respiration but this positive effect was cancelled out by the significantly negative effect of the increased soil total N concentration. The decrease in soil pH was the most effective pathway to decrease TBCF in high-N addition. Because of a large-scale reforestation program for C sink management in recent years, our findings of the nonlinear response of TBCF to different N fertilization treatments could provide insight for predicting belowground C sequestration potential and its response to atmospheric N deposition.

Inorganic N addition replaces N supplied to switchgrass (Panicum virgatum) by arbuscular mycorrhizal fungi.

Arbuscular mycorrhizal fungi (AMF) provide many benefits in agroecosystems including improved soil tilth, carbon sequestration, and water and nutrient transfer to plants. AMF are known to affect plant nitrogen (N) dynamics and transfer N to plants, but there have been few studies addressing whether the amount of N transferred to plants by AMF is agronomically relevant. We used δ15 N natural abundance methods and δ15 N mass balance equations to estimate the amount of plant N derived from AMF transfer in perennial grasses managed for bioenergy production under different N addition treatments (0, 56, and 196kg N/ha). Differentiation of δ15 N among plant, soil N, and AMF pools was higher than anticipated leading to calculations of 34-55% of plant N transferred by AMF in the treatments receiving no N addition to 6-22% of plant N transferred to plants in high-N addition treatments. AMF extra-radical hyphae biomass was significantly reduced in the high-N (196kg N/ha) addition treatments, which was negatively correlated to enriched plant δ15 N. Our results suggest that N addition decreases AMF N transfer to plants. When N was limiting to plant growth, AMF supplied agronomically significant amounts of plant N, and a higher proportion of overall plant N. Because differentiation between N pools was greater than expected, stable isotope measurements can be used to estimate N transfer to AMF plant hosts.

Long-term nitrogen addition modifies microbial composition and functions for slow carbon cycling and increased sequestration in tropical forest soil.

Nitrogen (N) deposition is a component of global change that has considerable impact on belowground carbon (C) dynamics. Plant growth stimulation and alterations of fungal community composition and functions are the main mechanisms driving soil C gains following N deposition in N-limited temperate forests. In N-rich tropical forests, however, N deposition generally has minor effects on plant growth; consequently, C storage in soil may strongly depend on the microbial processes that drive litter and soil organic matter decomposition. Here, we investigated how microbial functions in old-growth tropical forest soil responded to 13years of N addition at four rates: 0 (Control), 50 (Low-N), 100 (Medium-N), and 150 (High-N) kgNha-1 year-1 . Soil organic carbon (SOC) content increased under High-N, corresponding to a 33% decrease in CO2 efflux, and reductions in relative abundances of bacteria as well as genes responsible for cellulose and chitin degradation. A 113% increase in N2 O emission was positively correlated with soil acidification and an increase in the relative abundances of denitrification genes (narG and norB). Soil acidification induced by N addition decreased available P concentrations, and was associated with reductions in the relative abundance of phytase. The decreased relative abundance of bacteria and key functional gene groups for C degradation were related to slower SOC decomposition, indicating the key mechanisms driving SOC accumulation in the tropical forest soil subjected to High-N addition. However, changes in microbial functional groups associated with N and P cycling led to coincidentally large increases in N2 O emissions, and exacerbated soil P deficiency. These two factors partially offset the perceived beneficial effects of N addition on SOC storage in tropical forest soils. These findings suggest a potential to incorporate microbial community and functions into Earth system models considering their effects on greenhouse gas emission, biogeochemical processes, and biodiversity of tropical ecosystems.

Responses of soil respiration to nitrogen addition in the Sanjiang Plain wetland, northeastern China.

This study was designed to test the hypothesis that nitrogen (N) addition leads to enhanced soil respiration (SR) in nitrogen deficient marsh. Here, we report the response of SR to simulated N deposition in a temperate marsh of northeastern China from June 2009 to September 2011. The experiment included three-levels of N treatment (control: no N addition, Low-N: 4g N m-2 y-1, and High-N: 8 g N m-2 y-1). Our study showed various responses of SR to level and duration of N addition. Yearly SR was increased by 11.8%-15.2% (P<0.05) under Low-N addition during the three years, while SR showed a strong increase by 27.5% (P<0.05) in the first year and then decreased by 4.4% (P>0.05) and 15.4% (P<0.05) in the next two years under High-N addition. Soil respiration was positively correlated with soil temperature and negatively correlated with soil water content. High-N treatment reduced soil pH value (P<0.05). The negative response of SR to High-N addition in the following two years may attribute to lower microbial activity, microbial biomass and alteration in the microbial community due to lower soil pH, which consequently leads to decreased SR. Meanwhile, we found root biomass were increased under High-N addition. This implies that the increase of autotrophic respiration was lower than the decline of heterotrophic respiration in the following two years. Our findings suggest complex interactions between N deposition and SR, which is needed to be further investigated in the future studies.

Changes in litter quality induced by N deposition alter soil microbial communities

Soil microbial community composition and litter quality are important drivers of litter decomposition, but how litter quality influences the soil microbial composition largely remains unknown. We conducted a microcosm experiment to examine the effects of changes in litter quality induced by long-term N deposition on soil microbial community composition. Mixed-species litter and single-species litter were collected from a field experiment with replicate plots exposed to long-term N-addition in a semiarid grassland in northern China. The litters were decomposed in a standard live soil after which the composition of the microbial community was determined by Illumina MiSeq Sequencing. Changes in litter stoichiometry induced by N-addition increased the diversity of the fungal community. The alpha-diversity of the fungal community was more sensitive to the type of litter (mixed- or single-species) than to the N-addition effects, with higher abundance of fungal OTUs and Shannon-diversity observed in soil with mixed-species litter. Moreover, the relative abundance of saprophytic fungi increased with increasing N-addition rates, which suggests that fungi play an important role in the initial stages of the decomposition process. Litter type and N addition did not significantly change the diversity of bacterial community. The relative abundance of ammonia-oxidizing bacteria was lower in high N-addition treatments than in those with lower N input, indicating that changes in litter stoichiometry could change ecosystem functioning via its effects on bacteria. Our results presented robust evidence for the plant-mediated pathways through which N-deposition affects the soil microbial community and biogeochemical cycling.

Effects of litter manipulation on soil respiration under short-term nitrogen addition in a subtropical evergreen forest

Nitrogen (N) availability is rapidly increasing in subtropical ecosystems, where litterfall is also accelerating and may substantially affect belowground carbon (C) storage and soil respiration (Rs). This study aims to detect how litter inputs affect Rs under N addition in a subtropical forest. We conducted a two-factor experiment (N addition and litter manipulation) in a subtropical Schima superba evergreen broad-leaved forest in eastern China. Three levels of N addition included low-N (50 kg N ha−1 yr−1), high-N (100 kg N ha−1 yr−1) and ambient N (0 kg N ha−1 yr−1), and three levels of litter manipulation consisting of litter removal (NL), litter addition (DL) and Control litter input were conducted. Our results showed NL decreased Rs by 41% and 38% under low- and high-N additions, respectively, compared to Control litter input. DL decreased Rs by 24% compared to Control litter input under high-N addition. Furthermore, low- and high-N additions decreased the effect size of NL on Rs, whereas high-N addition decreased the effect size of DL on Rs in the study period, in both rainy and dry seasons. The effect of litter input alteration on Rs under N addition decreased, compared to Rs in Control litter input under N addition. Thus, the increase in Rs under N addition was alleviated, suggesting that soil organic C sequestration may benefit from increasing N deposition in the future. Further study is needed to clarify the effects of litter input alteration on Rs and soil C cycling under long-term N addition in subtropical forests.

Negative effect of nitrogen addition on soil respiration dependent on stand age: Evidence from a 7-year field study of larch plantations in northern China

Increasing anthropogenic nitrogen (N) deposition will affect the global carbon (C) cycle, although the mechanisms and magnitudes of its effects are not yet fully understood. Soil respiration (Rs) and its components (i.e., heterotrophic (Rh) and autotrophic (Ra) respiration) represent the largest flux of carbon dioxide (CO2) from terrestrial ecosystems to the atmosphere. However, the potentially interactive effect of N addition and stand age on Rs in forests remains unclear. We conducted a 7-year (2010–2016) field experiment in three differently-aged larch plantations (11-, 20-, and 45-year-old, representing sapling, intermediate, and mature stands) to investigate the effects of different N addition rates (control: no N addition; low-N addition, N20: 20 kg N ha−1 year−1; and high-N addition, N50: 50 kg N ha−1 year−1) on Rs. We found significant seasonal variation in Rs, Rh, and Ra, with levels highest in summer and lowest in autumn. Rs correlated strongly with soil temperature and weakly with soil moisture, and Rs, Rh, and Ra increased exponentially with soil temperature. The temperature sensitivity (Q10) of Rs was lowest in the mature stand, which suggests that it will sequester relatively more C in a warmer climate. The Q10 of Ra was higher than that of Rh, indicating that it contributed proportionally more to the total soil CO2 efflux. N addition had no significant effect on Q10. Rs in the intermediate stand was significantly lower than Rs in the sapling and mature stands, which may be due to its lower soil temperature and poorer substrate quality. N addition decreased Rs by an average of 9%, but significant effects were only observed in the sapling and intermediate stands. The negative influence of N addition on Rs was largely dependent on soil temperature, with more negative effects observed in the summer when temperatures were higher, and attributable to a reduction in either Ra in the intermediate stand or Rh in the sapling stand. We hypothesize that N limitation in the young trees was alleviated by fertilization, which decreased C allocation to roots, whereas N addition may have exerted a more negative influence on soil microbial activity in the sapling stand. In summary, our results highlight the importance of stand age in regulating the negative effects of N addition on Rs and its components, which has implications for projecting the global C–climate feedback in the future.

Sequestration of atmospheric CO2 in boreal forest carbon pools in northeastern China: Effects of nitrogen deposition

An increase in nitrogen (N) deposition has been proposed to cause boreal forests to capture and store a globally significant quantity of carbon (C), but the size of the boreal forest C sink remains uncertain after N addition. Therefore, we conducted a N addition experiment using four N addition rates (0, 2.5, 5.0 and 7.5gNm−2yr−1) in the boreal zone of northeastern China to determine the changes in forest C sequestration and to investigate the mechanisms of the changes in C sequestration after N addition. Our data show that N addition increases the total C sequestration, but the efficiency of this effect is reduced as the N addition rate increases. We also found that the amount and the mechanism of the C sequestration increase in above- and belowground C pools vary with different amounts of N addition. Low- and medium-N addition increased the above- and belowground C sequestration, and the potential mechanisms responsible for such C accumulation include N-induced increases in photosynthesis via a decrease in the foliar C content and increases in root mass via increased plant C allocation in the roots. However, high-N addition decreases aboveground C sequestration by inhibiting photosynthesis and increases belowground C sequestration by inhibiting soil C losses. Our data indicate that the response patterns of above- and belowground C pools to different amounts of N addition may involve several complex biochemical processes and occur by different mechanisms; therefore, separating the effects of N addition on above- and belowground C sequestration will help improve and validate current modeling efforts.