Higher aridity and more extreme rainfall events in drylands are predicted due to climate change. Yet, it is unclear how changing precipitation regimes may affect nitrogen (N) cycling, especially in areas with extremely high aridity. Here we investigate soil N isotopic values (d 15 N) along a 3,200 km aridity gradient and reveal a hump-shaped relationship between soil d 15 N and aridity index (AI) with a threshold at AI ¼ 0.32. Variations of foliar d 15 N, the abundance of nitrification and denitrification genes, and metabolic quotient along the gradient provide further evidence for the existence of this threshold. Data support the hypothesis that the increase of gaseous N loss is higher than the increase of net plant N accumulation with increasing AI below AI ¼ 0.32, while the opposite is favoured above this threshold. Our results highlight the importance of N-cycling microbes in extremely dry areas and suggest different controlling factors of N-cycling on either side of the threshold.

Ecosystem structure, functioning and stability have been a focus of ecological and environmental sciences during the past two decades. The mechanisms underlying their relationship, however, are not well understood. Based on comprehensive studies in Inner Mongolia grassland, here we show that species-level stoichiometric homoeostasis was consistently positively correlated with dominance and stability on both 2-year and 27-year temporal scales and across a 1200-km spatial transect. At the community level, stoichiometric homoeostasis was also positively correlated with ecosystem function and stability in most cases. Thus, homoeostatic species tend to have high and stable biomass; and ecosystems dominated by more homoeostatic species have higher productivity and greater stability. By modulating organism responses to key environmental drivers, stoichiometric homoeostasis appears to be a major mechanism responsible for the structure, functioning and stability of grassland ecosystems.

Atmospheric concentrations of the greenhouse gas nitrous oxide (N(2)O) have increased significantly since pre-industrial times owing to anthropogenic perturbation of the global nitrogen cycle, with animal production being one of the main contributors. Grasslands cover about 20 per cent of the temperate land surface of the Earth and are widely used as pasture. It has been suggested that high animal stocking rates and the resulting elevated nitrogen input increase N(2)O emissions. Internationally agreed methods to upscale the effect of increased livestock numbers on N(2)O emissions are based directly on per capita nitrogen inputs. However, measurements of grassland N(2)O fluxes are often performed over short time periods, with low time resolution and mostly during the growing season. In consequence, our understanding of the daily and seasonal dynamics of grassland N(2)O fluxes remains limited. Here we report year-round N(2)O flux measurements with high and low temporal resolution at ten steppe grassland sites in Inner Mongolia, China. We show that short-lived pulses of N(2)O emission during spring thaw dominate the annual N(2)O budget at our study sites. The N(2)O emission pulses are highest in ungrazed steppe and decrease with increasing stocking rate, suggesting that grazing decreases rather than increases N(2)O emissions. Our results show that the stimulatory effect of higher stocking rates on nitrogen cycling and, hence, on N(2)O emission is more than offset by the effects of a parallel reduction in microbial biomass, inorganic nitrogen production and wintertime water retention. By neglecting these freeze-thaw interactions, existing approaches may have systematically overestimated N(2)O emissions over the last century for semi-arid, cool temperate grasslands by up to 72 per cent.

Stoichiometric homeostasis, the degree to which an organism maintains its C:N:P ratios around a given species- or stage-specific value despite variation in the relative availabilities of elements in its resource supplies, is a key parameter in ecological stoichiometry. However, its regulation and role in affecting organismal and ecosystem processes is still poorly understood in vascular plants. We performed a sand culture experiment and a field nitrogen (N) and phosphorus (P) addition experiment to evaluate the strength of N, P and N:P homeostasis in higher plants in the Inner Mongolia grassland. Our results showed that homeostatic regulation coefficients (H) of vascular plants ranged from 1.93 to 14.5. H varied according to plant species, aboveground and belowground compartments, plant developmental stage, and overall plant nutrient content and N:P ratio. H for belowground and for foliage were inversely related, while H increased with plant developmental stage. H for N (H(N)) was consistently greater than H for P (H(P)) while H for N:P (H(N:P)) was consistently greater than H(N) and H(P). Furthermore, species with greater N and P contents and lower N:P were less homeostatic, suggesting that more homeostatic plants are more conservative nutrient users. The results demonstrate that H of plants encompasses a considerable range but is stronger than that of algae and fungi and weaker than that of animals. This is the first comprehensive evaluation of factors influencing stoichiometric homeostasis in vascular plants.

The growth rate hypothesis (GRH) proposes that higher growth rate (the rate of change in biomass per unit biomass, μ) is associated with higher P concentration and lower C∶P and N∶P ratios. However, the applicability of the GRH to vascular plants is not well-studied and few studies have been done on belowground biomass. Here we showed that, for aboveground, belowground and total biomass of three study species, μ was positively correlated with N∶C under N limitation and positively correlated with P∶C under P limitation. However, the N∶P ratio was a unimodal function of μ, increasing for small values of μ, reaching a maximum, and then decreasing. The range of variations in μ was positively correlated with variation in C∶N∶P stoichiometry. Furthermore, μ and C∶N∶P ranges for aboveground biomass were negatively correlated with those for belowground. Our results confirm the well-known association of growth rate with tissue concentration of the limiting nutrient and provide empirical support for recent theoretical formulations.

Plant carbon : nitrogen : phosphorus (C:N:P) ratios are powerful indicators of diverse ecological processes. During plant development and growth, plant C:N:P stoichiometry responds to environmental conditions and physiological constraints. However, variations caused by effects of sampling (i.e. sampling date, leaf age and root size) often have been neglected in previous studies. We investigated the relative contributions of sampling date, leaf age, root size and species identity to stoichiometric flexibility in a field mesocosm study and a natural grassland in Inner Mongolia. We found that sampling date, leaf age, root size and species identity all significantly affected C:N:P stoichiometry both in the pot study as well as in the field. Overall, C:N and C:P ratios increased significantly over time and with increasing leaf age and root size, while the dynamics of N:P ratios depended on species identity. Our results suggest that attempts to synthesize C:N:P stoichiometry data across studies that span regional to global scales and include many species need to better account for temporal variation.

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

Nitrogen (N) deposition has been steadily increasing for decades, with consequences for soil respiration. However, we have a limited understanding of how soil respiration responds to N availability. Here, we investigated the soil respiration responses to low and high levels of N addition (0.4 mol N m−2 yr−1 vs 1.6 mol N m−2 yr−1) over a two-year period in a semiarid Leymus chinensis grassland in Inner Mongolia, China. Our results show that low-level N addition increased soil respiration, plant belowground biomass and soil microbial biomass carbon (MBC), while high-level N additions decreased them. Soil respiration was positively correlated with plant belowground biomass, MBC, soil temperature and soil moisture. Together plant belowground biomass and MBC explained 99.4% of variation in mean soil respiration, with plant belowground biomass explaining 63.4% of the variation and soil MBC explaining the remaining 36%. Finally, the temperature sensitivity of soil respiration was not influenced by N additions. Overall, our results suggest that low levels of N deposition may stimulate soil respiration, but large increases in N availability may decrease soil respiration, and that these responses are driven by the dissimilar responses of both plant belowground biomass and soil MBC.