Global patterns and substrate‐based mechanisms of the terrestrial nitrogen cycle

Nitrogen (N) deposition is impacting the services that ecosystems provide to humanity. However, the mechanisms determining impacts on the N cycle are not fully understood. To explore the mechanistic underpinnings of N impacts on N cycle processes, we reviewed and synthesised recent progress in ecosystem N research through empirical studies, conceptual analysis and model simulations. Experimental and observational studies have revealed that the stimulation of plant N uptake and soil retention generally diminishes as N loading increases, while dissolved and gaseous losses of N occur at low N availability but increase exponentially and become the dominant fate of N at high loading rates. The original N saturation hypothesis emphasises sequential N saturation from plant uptake to soil retention before N losses occur. However, biogeochemical models that simulate simultaneous competition for soil N substrates by multiple processes match the observed patterns of N losses better than models based on sequential competition. To enable better prediction of terrestrial N cycle responses to N loading, we recommend that future research identifies the response functions of different N processes to substrate availability using manipulative experiments, and incorporates the measured N saturation response functions into conceptual, theoretical and quantitative analyses.     

Grazing intensity significantly affects belowground carbon and nitrogen cycling in grassland ecosystems: a meta‐analysis

Livestock grazing activities potentially alter ecosystem carbon (C) and nitrogen (N) cycles in grassland ecosystems. Despite the fact that numerous individual studies and a few meta‐analyses had been conducted, how grazing, especially its intensity, affects belowground C and N cycling in grasslands remains unclear. In this study, we performed a comprehensive meta‐analysis of 115 published studies to examine the responses of 19 variables associated with belowground C and N cycling to livestock grazing in global grasslands. Our results showed that, on average, grazing significantly decreased belowground C and N pools in grassland ecosystems, with the largest decreases in microbial biomass C and N (21.62% and 24.40%, respectively). In contrast, belowground fluxes, including soil respiration, soil net N mineralization and soil N nitrification increased by 4.25%, 34.67% and 25.87%, respectively, in grazed grasslands compared to ungrazed ones. More importantly, grazing intensity significantly affected the magnitude (even direction) of changes in the majority of the assessed belowground C and N pools and fluxes, and C : N ratio as well as soil moisture. Specifically,light grazing contributed to soil C and N sequestration whereas moderate and heavy grazing significantly increased C and N losses. In addition, soil depth, livestock type and climatic conditions influenced the responses of selected variables to livestock grazing to some degree. Our findings highlight the importance of the effects of grazing intensity on belowground C and N cycling, which may need to be incorporated into regional and global models for predicting effects of human disturbance on global grasslands and assessing the climate‐biosphere feedbacks.     

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.     

Effects of mild winter freezing on soil nitrogen and carbon dynamics in a northern hardwood forest

Overwinter and snowmelt processes are thought to be critical to controllersof nitrogen (N) cycling and retention in northern forests. However, therehave been few measurements of basic N cycle processes (e.g.mineralization, nitrification, denitrification) during winter and littleanalysis of the influence of winter climate on growing season N dynamics.In this study, we manipulated snow cover to assess the effects of soilfreezing on in situ rates of N mineralization, nitrification and soilrespiration, denitrification (intact core, C₂H₂ – based method),microbial biomass C and N content and potential net N mineralization andnitrification in two sugar maple and two yellow birch stands with referenceand snow manipulation treatment plots over a two year period at theHubbard Brook Experimental Forest, New Hampshire, U.S.A. The snowmanipulation treatment, which simulated the late development of snowpackas may occur in a warmer climate, induced mild (temperatures >–5 °C) soil freezing that lasted until snowmelt. The treatmentcaused significant increases in soil nitrate (NO₃ ^–)concentrations in sugar maple stands, but did not affect mineralization,nitrification, denitrification or microbial biomass, and had no significanteffects in yellow birch stands. Annual N mineralization and nitrificationrates varied significantly from year to year. Net mineralization increasedfrom 12.0 g N m^–2 y^–1 in 1998 to 22 g N m^–2 y^–1 in 1999 and nitrification increased from 8 g N m^–2 y^–1 in 1998 to 13 g N m^–2 y^–1 in 1999.Denitrification rates ranged from 0 to 0.65 g N m^–2 y^–1. Ourresults suggest that mild soil freezing must increase soil NO₃ ^– levels by physical disruption of the soil ecosystem and not by direct stimulation of mineralization and nitrification. Physical disruption canincrease fine root mortality, reduce plant N uptake and reduce competitionfor inorganic N, allowing soil NO₃ ^– levels to increase evenwith no increase in net mineralization or nitrification.     

Responses of plant phenology to nitrogen addition: a meta‐analysis

Phenology is one of the most sensitive processes of plant in response to global change. Anthropogenic activities have considerably increased nitrogen (N) deposition, which significant affects plant phenology. Although numerous individual studies have been conducted, it remains controversial how N addition affects phenological stages, and a comprehensive understanding of how plant phenology responds to external N inputs remains elusive. To reconcile the differences, we conducted a meta‐analysis of 117 species to examine the responses of plant phenology to N addition in terrestrial ecosystems, and assessed variations in their responses in relation to ecosystem types, functional groups, and environmental conditions. Our results showed that plant phenology changed significantly after N addition, and phenology time delayed and phenology duration shortened significantly across all biomes except fruiting duration, but varied with biome types. The phenology change in cropland was more dramatical than in grassland after N addition, even in opposite directions. The response of phenological stages to N addition was consistent in two pollination types except the flowering time, the flowering time had no change in anemophilous but significantly delayed in entomophilous. In addition, the response of phenology to N addition was discrepancy among functional groups, the phenology time advanced and duration shortened in sedge, while phenology time delayed and duration shortened in other groups, and the phenology change in legume was larger than grass and forbs. We also found that environmental factors had little effects on the response of plant phenology to N addition, but significant correlation was found between the response ratios of different phenological stages. Our study suggested that phenology was sensitive to N deposition at many phenological stages, and changes in phenology may be smaller with community biodiversity increasing at ecosystem level.     

Responses of terrestrial ecosystems to temperature and precipitation change: a meta‐analysis of experimental manipulation

Global mean temperature is predicted to increase by 2–7 °C and precipitation to change across the globe by the end of this century. To quantify climate effects on ecosystem processes, a number of climate change experiments have been established around the world in various ecosystems. Despite these efforts, general responses of terrestrial ecosystems to changes in temperature and precipitation, and especially to their combined effects, remain unclear. We used meta‐analysis to synthesize ecosystem‐level responses to warming, altered precipitation, and their combination. We focused on plant growth and ecosystem carbon (C) balance, including biomass, net primary production (NPP), respiration, net ecosystem exchange (NEE), and ecosystem photosynthesis, synthesizing results from 85 studies. We found that experimental warming and increased precipitation generally stimulated plant growth and ecosystem C fluxes, whereas decreased precipitation had the opposite effects. For example, warming significantly stimulated total NPP, increased ecosystem photosynthesis, and ecosystem respiration. Experimentally reduced precipitation suppressed aboveground NPP (ANPP) and NEE, whereas supplemental precipitation enhanced ANPP and NEE. Plant productivity and ecosystem C fluxes generally showed higher sensitivities to increased precipitation than to decreased precipitation. Interactive effects of warming and altered precipitation tended to be smaller than expected from additive, single‐factor effects, though low statistical power limits the strength of these conclusions. New experiments with combined temperature and precipitation manipulations are needed to conclusively determine the importance of temperature–precipitation interactions on the C balance of terrestrial ecosystems under future climate conditions.     

Global response patterns of plant photosynthesis to nitrogen addition: A meta‐analysis

A mechanistic understanding of plant photosynthetic response is needed to reliably predict changes in terrestrial carbon (C) gain under conditions of chronically elevated atmospheric nitrogen (N) deposition. Here, using 2,683 observations from 240 journal articles, we conducted a global meta‐analysis to reveal effects of N addition on 14 photosynthesis‐related traits and affecting moderators. We found that across 320 terrestrial plant species, leaf N was enhanced comparably on mass basis (N_mass, +18.4%) and area basis (N_area, +14.3%), with no changes in specific leaf area or leaf mass per area. Total leaf area (TLA) was increased significantly, as indicated by the increases in total leaf biomass (+46.5%), leaf area per plant (+29.7%), and leaf area index (LAI, +24.4%). To a lesser extent than for TLA, N addition significantly enhanced leaf photosynthetic rate per area (A_area, +12.6%), stomatal conductance (g_s, +7.5%), and transpiration rate (E, +10.5%). The responses of A_area were positively related with that of g_s, with no changes in instantaneous water‐use efficiency and only slight increases in long‐term water‐use efficiency (+2.5%) inferred from ¹³C composition. The responses of traits depended on biological, experimental, and environmental moderators. As experimental duration and N load increased, the responses of LAI and A_area diminished while that of E increased significantly. The observed patterns of increases in both TLA and E indicate that N deposition will increase the amount of water used by plants. Taken together, N deposition will enhance gross photosynthetic C gain of the terrestrial plants while increasing their water loss to the atmosphere, but the effects on C gain might diminish over time and that on plant water use would be amplified if N deposition persists.     

Nitrogen dynamics in grain crop and legume pasture systems under elevated atmospheric carbon dioxide concentration: A meta‐analysis

Understanding nitrogen (N) removal and replenishment is crucial to crop sustainability under rising atmospheric carbon dioxide concentration ([CO₂]). While a significant portion of N is removed in grains, the soil N taken from agroecosystems can be replenished by fertilizer application and N₂ fixation by legumes. The effects of elevated [CO₂] on N dynamics in grain crop and legume pasture systems were evaluated using meta‐analytic techniques (366 observations from 127 studies). The information analysed for non‐legume crops included grain N removal, residue C : N ratio, fertilizer N recovery and nitrous oxide (N₂O) emission. In addition to these parameters, nodule number and mass, nitrogenase activity, the percentage and amount of N fixed from the atmosphere were also assessed in legumes. Elevated [CO₂] increased grain N removal of C₃ non‐legumes (11%), legumes (36%) and C₄ crops (14%). The C : N ratio of residues from C₃ non‐legumes and legumes increased under elevated [CO₂] by 16% and 8%, respectively, but the increase for C₄ crops (9%) was not statistically significant. Under elevated [CO₂], there was a 38% increase in the amount of N fixed from the atmosphere by legumes, which was accompanied by greater whole plant nodule number (33%), nodule mass (39%), nitrogenase activity (37%) and %N derived from the atmosphere (10%; non‐significant). Elevated [CO₂] increased the plant uptake of fertilizer N by 17%, and N₂O emission by 27%. These results suggest that N demand and removal in grain cropping systems will increase under future CO₂‐enriched environments, and that current N management practices (fertilizer application and legume incorporation) will need to be revised.     

不同氮输入对湿地草甸沼泽土N_2O排放和有机碳矿化的影响

通过室内培养实验,研究了不同氮输入梯度下(N0:0mg·g-1,N1:0.1mg·g-1,N2:0.2mg·g-1,N3:0.5mg·g-1)湿地草甸沼泽土N2O排放和有机碳矿化特征,并分析了土壤微生物量碳、氮变化规律。整个培养期(23d)内,N0、N1、N2和N3处理N2O排放总量分别为91.12、133.02、147.75和303.45μg.kg-1,随氮输入量增大而增大,表明氮输入对N2O排放产生促进作用;氮输入处理的有机碳矿化速率在整个培养期除最后培养阶段外均低于对照,表明氮输入对有机碳矿化有一定的抑制作用;各氮输入处理土壤微生物量碳降低,与对照差异显著(P0.05),但各处理间差异未达到显著水平,土壤微生物量氮随氮输入量增大呈线性增加,各处理间差异显著(P0.05),表明氮输入影响土壤微生物结构和组成,具体影响机理须进一步探讨。     

Climate change amplifies gross nitrogen turnover in montane grasslands of Central Europe in both summer and winter seasons

The carbon‐ and nitrogen‐rich soils of montane grasslands are exposed to above‐average warming and to altered precipitation patterns as a result of global change. To investigate the consequences of climatic change for soil nitrogen turnover, we translocated intact plant–soil mesocosms along an elevational gradient, resulting in an increase of the mean annual temperature by approx. 2 °C while decreasing precipitation from approx. 1500 to 1000 mm. Following three years of equilibration, we monitored the dynamics of gross nitrogen turnover and ammonia‐oxidizing bacteria (AOB) and archaea (AOA) in soils over an entire year. Gross nitrogen turnover and gene levels of AOB and AOA showed pronounced seasonal dynamics. Both summer and winter periods equally contributed to cumulative annual N turnover. However, highest gross N turnover and abundance of ammonia oxidizers were observed in frozen soil of the climate change site, likely due to physical liberation of organic substrates and their rapid turnover in the unfrozen soil water film. This effect was not observed at the control site, where soil freezing did not occur due to a significant insulating snowpack. Climate change conditions accelerated gross nitrogen mineralization by 250% on average. Increased N mineralization significantly stimulated gross nitrification by AOB rather than by AOA. However, climate change impacts were restricted to the 2–6 cm topsoil and rarely occurred at 12–16 cm depth, where generally much lower N turnover was observed. Our study shows that significant mineralization pulses occur under changing climate, which is likely to result in soil organic matter losses with their associated negative impacts on key soil functions. We also show that N cycling processes in frozen soil can be hot moments for N turnover and thus are of paramount importance for understanding seasonal patterns, annual sum of N turnover and possible climate change feedbacks.     

High arctic heath soil respiration and biogeochemical dynamics during summer and autumn freeze‐in – effects of long‐term enhanced water and nutrient supply

In High Arctic NE Greenland, temperature and precipitation are predicted to increase during this century, however, relatively little information is available on the role of increased water supply on soil CO ₂ efflux in dry, high arctic ecosystems. We measured soil respiration (R_soil) in summer and autumn of 2009 in combination with microbial biomass and nutrient availability during autumn freeze‐in at a dry, open heath in Zackenberg, NE Greenland. This tundra site has been subject to fully factorial manipulation consisting of increased soil water supply for 14 years, and occasional nitrogen (N) addition in pulses. Summer watering enhanced R_soil during summer, but decreased R_soil in the following autumn. We speculate that this is due to intensified depletion of recently fixed plant carbon by soil organisms. Hence, autumn soil microbial activity seems tightly linked to growing season plant production through plant‐associated carbon pools. Nitrogen addition alone consistently increased R_soil, but when water and nitrogen were added in combination, autumn R_soil declined similarly to when water was added alone. Despite several freeze‐thaw events, the microbial biomass carbon (C) remained constant until finally being reduced by ~60% in late September. In spite of significantly reduced microbial biomass C and phosphorus (P), microbial N did not change. This suggests N released from dead microbes was quickly assimilated by surviving microbes. We observed no change in soil organic matter content after 14 years of environmental manipulations, suggesting high ecosystem resistance to environmental changes.     

Global‐scale impacts of nitrogen deposition on tree carbon sequestration in tropical,temperate, and boreal forests: A meta‐analysis

Elevated nitrogen (N) deposition may increase net primary productivity in N‐limited terrestrial ecosystems and thus enhance the terrestrial carbon (C) sink. To assess the magnitude of this N‐induced C sink, we performed a meta‐analysis on data from forest fertilization experiments to estimate N‐induced C sequestration in aboveground tree woody biomass, a stable C pool with long turnover times. Our results show that boreal and temperate forests responded strongly to N addition and sequestered on average an additional 14 and 13 kg C per kg N in aboveground woody biomass, respectively. Tropical forests, however, did not respond significantly to N addition. The common hypothesis that tropical forests do not respond to N because they are phosphorus‐limited could not be confirmed, as we found no significant response to phosphorus addition in tropical forests. Across climate zones, we found that young forests responded more strongly to N addition, which is important as many previous meta‐analyses of N addition experiments rely heavily on data from experiments on seedlings and young trees. Furthermore, the C–N response (defined as additional mass unit of C sequestered per additional mass unit of N addition) was affected by forest productivity, experimental N addition rate, and rate of ambient N deposition. The estimated C–N responses from our meta‐analysis were generally lower that those derived with stoichiometric scaling, dynamic global vegetation models, and forest growth inventories along N deposition gradients. We estimated N‐induced global C sequestration in tree aboveground woody biomass by multiplying the C–N responses obtained from the meta‐analysis with N deposition estimates per biome. We thus derived an N‐induced global C sink of about 177 (112–243) Tg C/year in aboveground and belowground woody biomass, which would account for about 12% of the forest biomass C sink (1,400 Tg C/year).     

Soil [N] modulates soil C cycling in CO2‐fumigated tree stands: a meta‐analysis

Under elevated atmospheric CO₂ concentrations, soil carbon (C) inputs are typically enhanced, suggesting larger soil C sequestration potential. However, soil C losses also increase and progressive nitrogen (N) limitation to plant growth may reduce the CO₂ effect on soil C inputs with time. We compiled a data set from 131 manipulation experiments, and used meta‐analysis to test the hypotheses that: (1) elevated atmospheric CO₂ stimulates soil C inputs more than C losses, resulting in increasing soil C stocks; and (2) that these responses are modulated by N. Our results confirm that elevated CO₂ induces a C allocation shift towards below‐ground biomass compartments. However, the increased soil C inputs were offset by increased heterotrophic respiration (Rh), such that soil C content was not affected by elevated CO₂. Soil N concentration strongly interacted with CO₂ fumigation: the effect of elevated CO₂ on fine root biomass and –production and on microbial activity increased with increasing soil N concentration, while the effect on soil C content decreased with increasing soil N concentration. These results suggest that both plant growth and microbial activity responses to elevated CO₂ are modulated by N availability, and that it is essential to account for soil N concentration in C cycling analyses.     

Drought effect on plant biomass allocation: A meta‐analysis

Drought is one of the abiotic stresses controlling plant function and ecological stability. In the context of climate change, drought is predicted to occur more frequently in the future. Despite numerous attempts to clarify the overall effects of drought stress on the growth and physiological processes of plants, a comprehensive evaluation on the impacts of drought stress on biomass allocation, especially on reproductive tissues, remains elusive. We conducted a meta‐analysis by synthesizing 164 published studies to elucidate patterns of plant biomass allocation in relation to drought stress. Results showed that drought significantly increased the fraction of root mass but decreased that of stem, leaf, and reproductive mass. Roots of herbaceous plants were more sensitive to drought than woody plants that reduced reproductive allocation more sharply than the former. Relative to herbaceous plants, drought had a more negative impact on leaf mass fraction of woody plants. Among the herbaceous plants, roots of annuals responded to drought stress more strongly than perennial herbs, but their reproductive allocation was less sensitive to drought than the perennial herbs. In addition, cultivated and wild plants seemed to respond to drought stress in a similar way. Drought stress did not change the scaling exponents of the allometric relationship between different plant tissues. These findings suggest that the allometric partitioning theory, rather than the optimal partitioning theory, better explains the drought‐induced changes in biomass allocation strategies.     

Aggravated phosphorus limitation on biomass production under increasing nitrogen loading: a meta‐analysis

Nitrogen (N) and phosphorus (P), either individually or in combination, have been demonstrated to limit biomass production in terrestrial ecosystems. Field studies have been extensively synthesized to assess global patterns of N impacts on terrestrial ecosystem processes. However, to our knowledge, no synthesis has been done so far to reveal global patterns of P impacts on terrestrial ecosystems, especially under different nitrogen (N) levels. Here, we conducted a meta‐analysis of impacts of P addition, either alone or with N addition, on aboveground (AGB) and belowground biomass production (BGB), plant and soil P concentrations, and N : P ratio in terrestrial ecosystems. Overall, our meta‐analysis quantitatively confirmed existing notions: (i) colimitation of N and P on biomass production and (ii) more P limitation in tropical forest than other ecosystems. More importantly, our analysis revealed new findings: (i) P limitation on biomass production was aggravated by N enrichment and (ii) plant P concentration was a better indicator of P limitation than soil P availability. Specifically, P addition increased AGB and BGB by 34% and 13%, respectively. The effect size of P addition on biomass production was larger in tropical forest than grassland, wetland, and tundra and varied with P fertilizer forms, P addition rates, or experimental durations. The P‐induced increase in biomass production and plant P concentration was larger under elevated than ambient N. Our findings suggest that the global limitation of P on biomass production will become severer under increasing N fertilizer and deposition in the future.