Soil carbon (C) and nitrogen (N) stoichiometry is a main driver of ecosystem functioning. Global N enrichment has greatly changed soil C : N ratios, but how altered resource stoichiometry influences the complexity of direct and indirect interactions among plants, soils, and microbial communities has rarely been explored. Here, we investigated the responses of the plant-soil-microbe system to multi-level N additions and the role of dissolved organic carbon (DOC) and inorganic N stoichiometry in regulating microbial biomass in semiarid grassland in northern China. We documented a significant positive correlation between DOC and inorganic N across the N addition gradient, which contradicts the negative nonlinear correlation between nitrate accrual and DOC availability commonly observed in natural ecosystems. Using hierarchical structural equation modeling, we found that soil acidification resulting from N addition, rather than changes in the plant community, was most closely related to shifts in soil microbial community composition and decline of microbial respiration. These findings indicate a down-regulating effect of high N availability on plant-microbe interactions. That is, with the limiting factor for microbial biomass shifting from resource stoichiometry to soil acidity, N enrichment weakens the bottom-up control of soil microorganisms by plant-derived C sources. These results highlight the importance of integratively studying the plant-soil-microbe system in improving our understanding of ecosystem functioning under conditions of global N enrichment.

Nitrogen (N) enrichment resulting from anthropogenic activities has greatly changed the composition and functioning of soil communities. Nematodes are one of the most abundant and diverse groups of soil organisms, and they occupy key trophic positions in the soil detritus food web. Nematodes have therefore been proposed as useful indicators for shifts in soil ecosystem functioning under N enrichment. Here, we monitored temporal dynamics of the soil nematode community using a multi-level N addition experiment in an Inner Mongolia grassland. Measurements were made three years after the start of the experiment. We used structural equation modeling (SEM) to explore the mechanisms regulating nematode responses to N enrichment. Across the N enrichment gradient, significant reductions in total nematode abundance, diversity (H' and taxonomic richness), maturity index (MI), and the abundance of root herbivores, fungivores and omnivores-predators were found in August. Root herbivores recovered in September, contributing to the temporal variation of total nematode abundance across the N gradient. Bacterivores showed a hump-shaped relationship with N addition rate, both in August and September. Ammonium concentration was negatively correlated with the abundance of total and herbivorous nematodes in August, but not in September. Ammonium suppression explained 61% of the variation in nematode richness and 43% of the variation in nematode trophic group composition. Ammonium toxicity may occur when herbivorous nematodes feed on root fluid, providing a possible explanation for the negative relationship between herbivorous nematodes and ammonium concentration in August. We found a significantly positive relationship between fungivores and fungal phospholipid fatty acids (PLFA), suggesting bottom-up control of fungivores. No such relationship was found between bacterivorous nematodes and bacterial PLFA. Our findings contribute to the understanding of effects of N enrichment in semiarid grassland on soil nematode trophic groups, and the cascading effects in the detrital soil food web.

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.

Higher ecosystem nitrogen (N) inputs resulting from human activities often suppress soil microbial biomass and respiration, thereby altering biogeochemical cycling. Soil acidification and carbon (C) limitation may drive these microbial responses, yet their relative importance remains elusive, which limits our understanding of the longer term effects of increasing N inputs. In a field experiment with continuous N addition at seven different rates from 0 to 50 g N m−2 year−1 over 6 years in a temperate grassland of Inner Mongolia, China, we examined the responses of soil microbial biomass and respiration to changes in soil acidity and C availability by adding lime and/or glucose to soil samples. Soil microbial biomass and respiration did only weakly respond to increasing soil pH, but increased strongly in response to higher C availability with increasing N addition rates. Soil net N immobilization increased in response to glucose addition, and soil microbial biomass increased at higher rates than microbial respiration along the gradient of previous N addition rates, both suggesting increasingly reinforced microbial C limitation with increasing N addition. Our results provide clear evidence for strong N‐induced microbial C limitation, but only little support for soil acidity effects within the initial pH range of 4.73–7.86 covered by our study. Field data support this conclusion by showing reduced plant C allocation belowground in response to N addition, resulting in soil microbial C starvation over the long term. In conclusion, soil microbial biomass and respiration under N addition were strongly dependent on C availability, most likely originating from plant belowground C inputs, and was much less affected by changes in soil pH. Our data help clarify a long‐standing debate about how increasing N input rates affect soil microbial biomass and respiration, and improve the mechanistic understanding of the linkages between ecosystem N enrichment and C cycling.

Grassland ecosystems cover c. 40% of global land area and contain c. 40% of soil organic carbon. Understanding the effects of adding nutrients to grasslands is essential because they provide much of our food, support diverse ecosystem services and harbor rich biodiversity.Using the meadow steppe (grassland) study site of Inner Mongolia, we manipulated seven key nutrients and a cocktail of micronutrients to examine their effects on grassland biomass productivity and diversity.The results, explained in structural equation models, link two previously disparate hypotheses in grassland ecology: (1) the light asymmetry competition hypothesis and (2) the genome size-nutrient interaction hypothesis. We show that aboveground net primary productivity increases predominantly from species with large genome sizes with the addition of nitrogen, and nitrogen plus phosphorus. This drives an asymmetric competition for light, causing a decline in species richness mainly in species with small genome sizes.This dynamic is likely to be caused by the nutrient demands of the nucleus and/or the scaling effects of nuclear size on cell size which impact water use efficiency. The model will help inform the best management approaches to reverse the rapid and unprecedented degradation of grasslands globally.

Phosphorus (P) limitation is expected to increase due to nitrogen (N)‐induced terrestrial eutrophication, although most soils contain large P pools immobilized in minerals (Pi) and organic matter (Po). Here we assessed whether transformations of these P pools could increase plant available pools alleviating P limitation under enhanced N availability. The mechanisms underlying these possible transformations were explored by combining results from a 10‐year field N addition experiment and a 3700‐km transect covering wide ranges in soil pH, soil N, aridity, leaching, and weathering that could affect soil P status in grasslands. Nitrogen addition promoted the dissolution of immobile Pi (mainly Ca‐bound recalcitrant P) to more available forms of Pi (including Al‐ and Fe‐bound P fractions and Olsen P) by decreasing soil pH from 7.6 to 4.7, but did not affect Po. Soil total P declined by 10% from 385 ± 6.8 to 346 ± 9.5 mg kg−1, whereas available P increased by 546% from 3.5 ± 0.3 to 22.6 ± 2.4 mg kg−1 after the 10‐year N addition, associated with an increase in Pi mobilization, plant uptake, and leaching. Similar to the N addition experiment, the drop in soil pH from 7.5 to 5.6 and increase in soil N concentration along the grassland transect were associated with an increased ratio between relatively mobile Pi and immobile Pi. Our results provide a new mechanistic understanding of the important role of soil Pi mobilization in maintaining plant P supply and accelerating biogeochemical P cycles under anthropogenic N enrichment. This mobilization process temporarily buffers ecosystem P limitation or even causes P eutrophication, but will extensively deplete soil P pools in the long run.

Temperature sensitivity (Q 10 ) of soil organic matter (SOM) decomposition is an important parameter in models of the global carbon (C) cycle. Previous studies have suggested that substrate quality controls the intrinsic Q 10 , whereas environmental factors can impose large constraints. For example, physical protection of SOM and its association with minerals attenuate the apparent Q 10 through reducing substrate availability and accessibility ([S]). The magnitude of this dampening effect, however, has never been quantified. We simulated theoretical Q 10 changes across a wide range of [S] and found that the relationship between Q 10 and the log 10 -transformed [S] followed a logistic rather than a linear function. Based on the unique Holocene paleosol chronosequence (7 soils from ca. 500 to 6900 years old), we demonstrated that the Q 10 decreased nonlinearly with soil age up to 1150 years, beyond which Q 10 remained stable. Hierarchical partitioning analysis indicated that an integrated C availability index, derived from principal component analysis of DOC content and parameters reflecting physical protection and mineral association, was the main explanatory variable for the nonlinear decrease of Q 10 with soil age. Microbial inoculation and 13 C-labelled glucose addition showed that low C availability induced by physical protection and minerals association attenuated Q 10 along the chronosequence. A separate soil incubation experiment indicated that Q 10 increased exponentially with activation energy (E a ) in the modern soil, suggesting that SOM chemical complexity regulates Q 10 only when C availability is high. In conclusion, organic matter availability strongly decreased with soil age, whereas Michelis-Menten kinetics defines the Q 10 response depending on | 4181 SU et al.

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