Different nitrogen saturation thresholds for above-, below-, and total net primary productivity in a temperate steppe.

Identifying the thresholds for the positive responses of total net primary productivity (NPP) to nitrogen (N) enrichment is an essential prerequisite for predicting the benefits of N deposition on ecosystem carbon sequestration. However, the responses of below-ground NPP (BNPP) to N enrichment are unknown in many ecosystems, which limits our ability to understand the carbon cycling under the scenario of increasing N availability. We examined the changes in above-ground NPP (ANPP), BNPP, and NPP of a temperate meadow steppe across a wide-ranging N addition gradient (0, 2, 5, 10, 20, and 50 g N m-2 year-1 ) during 5 years. Both ANPP and NPP increased nonlinearly with N addition rates. The N saturation threshold for ANPP (TA ) and NPP (TN ) was at the rate of 13.11 and 6.70 g N m-2 year-1 , respectively. BNPP decreased with increasing N addition when N addition rates ˃5 g N m-2 year-1 , resulting in much lower TN than TA . Soil N enrichment played a key role in driving the negative impacts of high N addition rates on BNPP, and consequently on the earlier occurrence of N saturation threshold for NPP. Our results highlight the negative effects of soil N enrichment on NPP in natural grasslands super-saturated with N. Furthermore, by considering ANPP and BNPP simultaneously, our results indicate that previous findings from above-ground might have over-estimated the positive effects of N deposition on primary productivity.

ß-diversity in temperate grasslands is driven by stronger environmental filtering of plant species with large genomes.

Elucidating mechanisms underlying community assembly and biodiversity patterns is central to ecology and evolution. Genome size (GS) has long been hypothesized to potentially affect species' capacity to tolerate environmental stress and might therefore help drive community assembly. However, its role in driving ß-diversity (i.e., spatial variability in species composition) remains unclear. We measured GS for 161 plant species and community composition across 52 sites spanning a 3200-km transect in the temperate grasslands of China. By correlating the turnover of species composition with environmental dissimilarity, we found that resource filtering (i.e., environmental dissimilarity that includes precipitation, and soil nitrogen and phosphorus concentrations) affected ß-diversity patterns of large-GS species more than small-GS species. By contrast, geographical distance explained more variation of ß-diversity for small-GS than for large-GS species. In a 10-year experiment manipulating levels of water, nitrogen, and phosphorus, adding resources increased plant biomass in species with large GS, suggesting that large-GS species are more sensitive to the changes in resource availability. These findings highlight the role of GS in driving community assembly and predicting species responses to global change.

Decoupled responses of above- and below-ground stability of productivity to nitrogen addition at the local and larger spatial scale.

Temporal stability of net primary productivity (NPP) is important for predicting the reliable provisioning of ecosystem services under global changes. Although nitrogen (N) addition is known to affect the temporal stability of aboveground net primary productivity (ANPP), it is unclear how it impacts that of belowground net primary productivity (BNPP) and NPP, and whether such effects are scale dependent. Here, using experimental N addition in a grassland, we found different responses of ANPP and BNPP stability to N addition at the local scale and that these responses propagated to the larger spatial scale. That is, N addition significantly decreased the stability of ANPP but did not affect the stability of BNPP and NPP at the two scales investigated. Additionally, spatial asynchrony of both ANPP and BNPP among communities provided greater stability at the larger scale and was not affected by N addition. Our findings challenge the traditional view that N addition would reduce ecosystem stability based on results from aboveground dynamics, thus highlighting the importance of viewing ecosystem stability from a whole system perspective.

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.

Effects of rhizoma peanut cultivars (Arachis glabrata Benth.) on the soil bacterial diversity and predicted function in nitrogen fixation.

There is a growing awareness of the importance of soil microorganisms in agricultural management practices. Currently, much less is known about whether different crop cultivar has an effect on the taxonomic structure and diversity, and specific functions of soil bacterial communities. Here, we examined the changes of the diversity and composition and enzyme-encoding nitrogenase genes in a long-term field experiment with seven different rhizoma peanut cultivars in southeastern USA, coupling high-throughput 16S rRNA gene sequencing and the sequence-based function prediction with Tax4Fun. Of the 32 phyla detected (Proteobacteria class), 13 were dominant: Acidobacteria, Alphaproteobacteria, Actinobacteria, Betaproteobacteria, Bacteroidetes, Verrucomicrobia, Gammaproteobacteria, Deltaproteobacteria, Gemmatimonadetes, Firmicutes, Nitrospirae, Chloroflexi, and Planctomycetes (relative abundance >1%). We found no evidence that the diversity and composition of bacterial communities were significantly different among different cultivars, but the abundance of some dominant bacterial groups that have N-fixation potentials (at broad or fine taxonomic level) and predicted abundances of some enzyme-encoding nitrogenase genes showed significant across-cultivar differences. The nitrogenase genes were notably abundant in Florigraze and Latitude soils while remarkably lower in Arbook and UF_TITO soils when compared with other cultivars, indicating different nitrogen fixation potentials among different cultivars. The findings also suggest that the abundance of certain bacterial taxa and the specific function bacteria perform in ecosystems can have an inherent association. Our study is helpful to understand how microbiological responses and feedback to different plant genotypes through the variation in structure and function of their communities in the rhizosphere.

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Plant genome size (GS) varies greatly over 2400-fold in angiosperms. Genome sizes are closely related to plant traits from cellular to individual level, which would have far-reaching ecolo-gical implications. Genome size may shape the interspecific responses of plants to changes of resource availability in Inner Mongolia grassland which is co-limited by water and nitrogen availabi-lity. We tested the role of genome size in structuring plant community composition after single and combined water (W) amd nitrogen (N) addition in a typical grassland of Inner Mongolia. Plant genome sizes were estimated by flow cytometry. We found that the response of plant aboveground net primary production (ANPP) to change in water availability was significantly affected by genome size. Water and NW addition significantly increased ANPP of small GS plants, instead of large GS species. Nitrogen addition had no effects on ANPP of both small and large GS plants. We found no effects of all the treatments on plant species richness. Results showed that GS modulated the response of grassland plant species to changes in water rather than nitrogen availability in Inner Mongolia. Since GS is a relatively constant trait with substantial interspecific variation, the application of GS in ecological studies would be of great significance to better understanding of ecosystem structure and function under global change.

Correction to: Mowing mitigates the negative impacts of N addition on plant species diversity.

Unfortunately, the panels of (f) in Figures 1, 2, and 4.

The relative contributions of intra- and inter-specific variation in driving community stoichiometric responses to nitrogen deposition and mowing in a grassland.

AIMS: The stoichiometric characteristics of plant communities are important controller for several fundamental ecological processes. The effects of environmental changes on community stoichiometric characteristics are driven by intra- and inter-specific variation. However, the relative importance of both pathways has seldom been empirically examined. METHODS: We quantified the relative contribution of intra- and inter-specific variation to the changes of community nitrogen (N) and phosphorus (P) concentrations after seven-year factorial N addition and mowing treatments in a semi-arid grassland of northern China. RESULTS: Nitrogen addition significantly increased community N and P concentrations and N:P ratio. Mowing significantly increased community N concentration and N:P. Intra-specific variation contributed more than inter-specific variation to the total variability of all the nutritional and stoichiometric characteristics, with intra-specific variation accounting for 68%, 70%, and 75% of the total variation in community-level N, P, and N:P, respectively. Negative covariations between the contribution of intra- and inter-specific variation occurred for community N and P concentrations. Further, N addition and mowing interacted to affect the impacts of intra- and inter-specific variation on community N concentration and N:P stoichiometry. CONCLUSIONS: Our results highlight different ways of trait selection for N addition and mowing treatments. Interactions between those two factors make it more difficult to accurately predict the responses of plant-mediated biogeochemical cycles under co-occurrence of environmental changes.

Mowing mitigates the negative impacts of N addition on plant species diversity.

Increasing availability of reactive nitrogen (N) threatens plant diversity in diverse ecosystems. While there is mounting evidence for the negative impacts of N deposition on one component of diversity, species richness, we know little about its effects on another one, species evenness. It is suspected that ecosystem management practice that removes nitrogen from the ecosystem, such as hay-harvesting by mowing in grasslands, would mitigate the negative impacts of N deposition on plant diversity. However, empirical evidence is scarce. Here, we reported the main and interactive effects of N deposition and mowing on plant diversity in a temperate meadow steppe with 4-year data from a field experiment within which multi-level N addition rates and multiple N compounds are considered. Across all the types of N compounds, species richness and evenness significantly decreased with the increases of N addition rate, which was mainly caused by the growth of a tall rhizomatous grass, Leymus chinensis. Such negative impacts of N addition were accumulating with time. Mowing significantly reduced the dominance of L. chinensis, and mitigated the negative impacts of N deposition on species evenness. We present robust evidence that N deposition threatened biodiversity by reducing both species richness and evenness, a process which could be alleviated by mowing. Our results highlight the changes of species evenness in driving the negative impacts of N deposition on plant diversity and the role of mowing in mediating such negative impacts of N deposition.

Environmental and spatial variables determine the taxonomic but not functional structure patterns of microbial communities in alpine grasslands.

There is considerable debate regarding how the taxonomic diversity of microbial communities relates to the functional diversity across space while similar questions have been explored in macro-organism communities. Here, we investigated the taxonomic and functional diversity patterns of soil microbial communities by coupling the data obtained from marker genes sequencing and functional gene surveys. Meanwhile, we evaluated the relative effects of environment and geographic distance on shaping these patterns in alpine grasslands of northern China. Although the taxonomic diversity and composition of microbial communities varied across sites, we found no consistent changes in the functional structure. Both the environmental factors and geographic distance concurrently affected the taxonomic diversity patterns but they had no effects on the spatial variations in functional genes. The functional alpha diversity was weakly correlated to the taxonomic alpha diversity across sites. Moreover, we found no significant relationship between the taxonomic and functional composition similarity among microbial communities. Together, our results provide evidence that spatial variation in microbial functions could be independent of their variations in taxonomic diversity. Even the drivers of spatial variations in the functional structure could be totally different from those of taxonomic variations such as environmental differences and dispersal limitation. Our findings suggest that spatial variations of microbial function structure within a community would not follow the variations of taxonomic structures due to different drivers between both of them over space.

The impacts of nitrogen deposition on community N:P stoichiometry do not depend on phosphorus availability in a temperate meadow steppe.

Nitrogen (N) enrichment has great consequences on several fundamental ecological processes through its impacts on plant nutrition traits (i.e. nutrient concentration and stoichiometric ratios); however, the extent to which the effects of N enrichment depend on phosphorus (P) availability are less well understood. While there is mounting evidence for the species-specific responses of plant nutrition traits to nutrient enrichment, we know little about the changes at the community-level. Here, we measured community-level biomass weighted (CWM) and non-weighted (CM) plant N and P concentrations and N:P ratio in a temperate meadow steppe after four years factorial N and P addition, with biomass and nutrition traits of each species in each plot being recorded. Nitrogen addition significantly increased community-level N concentration, decreased P concentration, and enhanced community N:P ratio. Phosphorus addition had no impacts on community-level N concentration, significantly increased P concentration, and reduced community N:P ratio. The impacts of N addition on community nutrition traits were not dependent on P addition and the community-level nutrition trait responses to N and P additions were primarily driven by intraspecific trait variation (ITV) rather than by species turnover. Community-level nutrition traits in the temperate meadow steppe were sensitive to the projected N and P enrichment. While nutrient enrichment had substantially changed community composition, its impacts on community nutrition traits were driven by ITV. Nitrogen deposition would result in imbalance of N and P in plant community, as indicated by the substantial increase in community-level N:P, which was not affected by increased P availability.

Dissolved methane in groundwater of domestic wells and its potential emissions in arid and semi-arid regions of Inner Mongolia, China.

Methane (CH4) is widely present in groundwater. Dissolved CH4 in groundwater is less understood when compared with that in wetlands. In this study, the concentrations and origin of dissolved CH4 in groundwater were investigated and the potential importance of groundwater CH4 emissions in arid and semi-arid regions of Inner Mongolia was discussed. Groundwater was extracted from domestic wells using a submersible pump or manual power and was analyzed for CH4 concentrations, δ13C-CH4, and physico-chemical variables. The results show that the concentrations of dissolved CH4 in groundwater had large spatial variability, ranging from 0 to 0.10 mg L-1 with a mean of 0.01 mg L-1 in Xilingol and from 0 to 8.99 mg L-1 with a mean of 1.44 mg L-1 in Xingan-Tongliao. Substantial CH4 concentrations of about 2.5-5.5 mg L-1 were found in central areas of Xingan-Tongliao in the winter and the summer. The δ13C-CH4 of about -85‰ was highly depleted while CH4 concentration was significantly negatively correlated with SO42- concentration, indicating that dissolved CH4 in groundwater was microbial in origin. This study suggests that groundwater as a source of CH4 might have great implications in arid and semi-arid regions worldwide and should deserve more research.

Carbon and nitrogen allocation shifts in plants and soils along aridity and fertility gradients in grasslands of China.

Plant carbon (C) and nitrogen (N) stoichiometry play an important role in the maintenance of ecosystem structure and function. To decipher the influence of changing environment on plant C and N stoichiometry at the subcontinental scale, we studied the shoot and root C and N stoichiometry in two widely distributed and dominant genera along a 2,200-km climatic gradient in China's grasslands. Relationships between C and N concentrations and soil climatic variables factors were studied. In contrast to previous theory, plant C concentration and C:N ratios in both shoots and roots increased with increasing soil fertility and decreased with increasing aridity. Relative N allocation shifted from soils to plants and from roots to shoots with increasing aridity. Changes in the C:N ratio were associated with changes in N concentration. Dynamics of plant C concentration and C:N ratios were mainly caused by biomass reallocation and a nutrient dilution effect in the plant-soil system. Our results suggest that the shifted allocation of C and N to different ecosystem compartments under a changing environment may change the overall use of these elements by the plant-soil system.

Habitat-specific patterns and drivers of bacterial ß-diversity in China's drylands.

The existence of biogeographic patterns among most free-living microbial taxa has been well established, yet little is known about the underlying mechanisms that shape these patterns. Here, we examined soil bacterial ß-diversity across different habitats in the drylands of northern China. We evaluated the relative importance of environmental factors versus geographic distance to a distance-decay relationship, which would be explained by the relative effect of basic ecological processes recognized as drivers of diversity patterns in macrobial theoretical models such as selection and dispersal. Although the similarity of bacterial communities significantly declined with increasing geographic distance, the distance-decay slope and the relative importance of factors driving distance-decay patterns varied across different habitats. A strong distance-decay relationship was observed in the alpine grassland, where the community similarity was influenced only by the environmental factors. In contrast, geographic distance was solely responsible for community similarity in the desert. Even the average compositional similarity among locations in the desert was distinctly lower compared with those in other habitats. We found no evidence that dispersal limitation strongly influenced the ß-diversity of bacterial communities in the desert grassland and typical grassland. Together, our results provide robust evidence of habitat specificity for microbial diversity patterns and their underlying drivers. Our findings suggest that microorganisms also have multiple drivers of diversity patterns and some of which may be parallel to some fundamental processes for explaining biodiversity patterns in macroorganisms.

Effects of mistletoe removal on growth, N and C reserves, and carbon and oxygen isotope composition in Scots pine hosts.

Most mistletoes are xylem-tapping hemiparasites, which derive their resources from the host's xylem solution. Thus, they affect the host's water relations and resource balance. To understand the physiological mechanisms underlying the mistletoe-host relationship, we experimentally removed Viscum album ssp. austriacum (Wiesb.) Vollmann from adult Pinus sylvestris L. host trees growing in a Swiss dry valley. We analyzed the effects of mistletoe removal over time on host tree growth and on concentrations of nonstructural carbohydrates (NSC) and nitrogen (N) in needles, fine roots and sapwood. In addition, we assessed the δ(13)C and δ(18)O in host tree rings. After mistletoe removal, δ(13)C did not change in newly produced tree rings compared with tree rings in control trees (still infected with mistletoe), but δ(18)O values increased. This pattern might be interpreted as a decrease in assimilation (A) and stomatal conductance (gs), but in our study, it most likely points to an inadequacy of the dual isotope approach. Instead, we interpret the unchanged δ(13)C in tree rings upon mistletoe removal as a balanced increase in A and gs that resulted in a constant intrinsic water use efficiency (defined as A/gs). Needle area-based concentrations of N, soluble sugars and NSC, as well as needle length, single needle area, tree ring width and shoot growth, were significantly higher in trees from which mistletoe was removed than in control trees. This finding suggests that mistletoe removal results in increased N availability and carbon gain, which in turn leads to increased growth rates of the hosts. Hence, in areas where mistletoe is common and the population is large, mistletoe management (e.g., removal) may be needed to improve the host vigor, growth rate and productivity, especially for relatively small trees and crop trees in xeric growth conditions.

Methane emissions from the trunks of living trees on upland soils.

Upland forests are traditionally thought to be net sinks for atmospheric methane (CH4 ). In such forests, in situ CH4 fluxes on tree trunks have been neglected relative to soil and canopy fluxes. We measured in situ CH4 fluxes from the trunks of living trees and other surfaces, such as twigs and soils, using a static closed-chamber method, and estimated the CH4 budget in a temperate upland forest in Beijing. We found that the trunks of Populus davidiana emitted large quantities of CH4 during July 2014-July 2015, amounting to mean annual emissions of 85.3 and 103.1 µg m(-2)  h(-1) on a trunk surface area basis on two replicate plots. The emission rates were similar in magnitude to those from tree trunks in wetland forests. The emitted CH4 was derived from the heartwood of trunks. On a plot or ecosystem scale, trunk CH4 emissions were equivalent to c. 30-90% of the amount of CH4 consumed by soils throughout the year, with an annual average of 63%. Our findings suggest that wet heartwoods, regardless of rot or not, occur widely in living trees on various habitats, where CH4 can be produced.

Impacts of leguminous shrub encroachment on neighboring grasses include transfer of fixed nitrogen.

Shrub encroachment induced by global change and human disturbance strongly affects ecosystem structure and function. In this study, we explore the degree to which invading leguminous shrubs affected neighboring grasses, including via the transfer of fixed nitrogen (N). We measured N concentrations and natural abundance (15)N of shoot tissues from three dominant grasses from different plant functional groups across seven distances along a local transect (up to 500 cm) to the leguminous shrub, Caragana microphylla. C. microphylla did transfer fixed N to neighboring grasses, but the amount and distance of N transferred were strongly species-specific. Shoot N concentrations decreased significantly with distance from C. microphylla, for a rhizomatous grass, Leymus chinensis, and a bunchgrass, Achnatherum sibiricum. However, N concentrations of another bunchgrass, Stipa grandis, were higher only directly underneath the shrub canopy. Shoot δ(15)N values of L. chinensis were enriched up to 500 cm from the shrub, but for S. grandis were enriched only below the shrub canopy. In contrast, δ(15)N of A. sibiricum did not change along the 500-cm transect. Our results indicated the rhizomatous grass transferred fixed N over long distances while bunchgrasses did not. The presence of C. microphylla increased the shoot biomass of L. chinensis but decreased that of S. grandis and A. sibiricum. These findings highlight the potential role of nutrient-acquisition strategies of neighboring grasses in moderating the interspecific variation of fixed N transfer from the leguminous shrub. Overall, leguminous shrubs have either positive or negative effects on the neighboring grasses and dramatically affect plant community composition and structure.

Plant nitrogen uptake drives responses of productivity to nitrogen and water addition in a grassland.

Increased atmospheric nitrogen (N) deposition and altered precipitation regimes have profound impacts on ecosystem functioning in semiarid grasslands. The interactions between those two factors remain largely unknown. A field experiment with N and water additions was conducted in a semiarid grassland in northern China. We examined the responses of aboveground net primary production (ANPP) and plant N use during two contrasting hydrological growing seasons. Nitrogen addition had no impact on ANPP, which may be accounted for by the offset between enhanced plant N uptake and decreased plant nitrogen use efficiency (NUE). Water addition significantly enhanced ANPP, which was largely due to enhanced plant aboveground N uptake. Nitrogen and water additions significantly interacted to affect ANPP, plant N uptake and N concentrations at the community level. Our observations highlight the important role of plant N uptake and use in mediating the effects of N and water addition on ANPP.

Effects of nitrogen addition and fire on plant nitrogen use in a temperate steppe.

Plant nitrogen (N) use strategies have great implications for primary production and ecosystem nutrient cycling. Given the increasing atmospheric N deposition received by most of the terrestrial ecosystems, understanding the responses of plant N use would facilitate the projection of plant-mediated N cycling under global change scenarios. The effects of N deposition on plant N use would be affected by both natural and anthropogenic disturbances, such as prescribed fire in the grassland. We examined the effects of N addition (5.25 g N m(-2) yr(-1)) and prescribed fire (annual burning) on plant N concentrations and N use characters at both species and community levels in a temperate steppe of northern China. We found that N addition and fire independently affected soil N availability and plant N use traits. Nitrogen addition increased aboveground net primary productivity (ANPP), inorganic N, and N uptake, decreased N response efficiency (NRE), but did not affect biomass-weighed N concentrations at community level. Prescribed fire did not change the community level N concentrations, but largely decreased N uptake efficiency and NRE. At the species level, the effects of N addition and fire on plant N use were species-specific. The divergent responses of plant N use at community and species levels to N addition and fire highlight the importance of the hierarchical responses of plant N use at diverse biological organization levels to the alteration of soil N availability. This study will improve our understanding of the responses of plant-mediated N cycling to global change factors and ecosystem management strategies in the semiarid grasslands.

Convergent responses of nitrogen and phosphorus resorption to nitrogen inputs in a semiarid grassland.

Human activities have significantly altered nitrogen (N) availability in most terrestrial ecosystems, with consequences for community composition and ecosystem functioning. Although studies of how changes in N availability affect biodiversity and community composition are relatively common, much less remains known about the effects of N inputs on the coupled biogeochemical cycling of N and phosphorus (P), and still fewer data exist regarding how increased N inputs affect the internal cycling of these two elements in plants. Nutrient resorption is an important driver of plant nutrient economies and of the quality of litter plants produce. Accordingly, resorption patterns have marked ecological implications for plant population and community fitness, as well as for ecosystem nutrient cycling. In a semiarid grassland in northern China, we studied the effects of a wide range of N inputs on foliar nutrient resorption of two dominant grasses, Leymus chinensis and Stipa grandis. After 4 years of treatments, N and P availability in soil and N and P concentrations in green and senesced grass leaves increased with increasing rates of N addition. Foliar N and P resorption significantly decreased along the N addition gradient, implying a resorption-mediated, positive plant-soil feedback induced by N inputs. Furthermore, N : P resorption ratios were negatively correlated with the rates of N addition, indicating the sensitivity of plant N and P stoichiometry to N inputs. Taken together, the results demonstrate that N additions accelerate ecosystem uptake and turnover of both N and P in the temperate steppe and that N and P cycles are coupled in dynamic ways. The convergence of N and P resorption in response to N inputs emphasizes the importance of nutrient resorption as a pathway by which plants and ecosystems adjust in the face of increasing N availability.