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Anthropogenic nitrogen (N) input is known to alter the soil microbiome, but how N enrichment influences the abundance, alpha-diversity and community structure of N-cycling functional microbial communities in grasslands remains poorly understood. Here, we collected soils from plant communities subjected to up to 9 years of annual N-addition (10 g N m −2 per year using urea as a N-source) and from unfertilized plots (control) in 30 grasslands worldwide spanning a large range of climatic and soil conditions. We focused on three key microbial groups responsible for two essential processes of the global N cycle: N 2 fixation (soil diazotrophs) and nitrification (AOA: ammonia-oxidizing archaea and AOB: ammonia-oxidizing bacteria). We targeted soil diazotrophs, AOA and AOB using Illumina MiSeq sequencing and measured the abundance (gene copy numbers) using quantitative PCR. N-addition shifted the structure of the diazotrophic communities, although their alpha-diversity and abundance were not affected. AOA and AOB responded differently to N-addition. The abundance and alpha-diversity of AOB increased, and their community structure shifted with N-addition. In contrast, AOA were not affected by N-addition. AOA abundance outnumbered AOB in control plots under conditions of low N availability, whereas N-addition favoured copiotrophic AOB. Overall, N-addition showed a low impact on soil diazotrophs and AOA while effects for AOB communities were considerable. These results reveal that long-term N-addition has important ecological implications for key microbial groups involved in two critical soil N-cycling processes. Increased AOB abundance and community shifts following N-addition may change soil N-cycling, as larger population sizes may promote higher rates of ammonia oxidation and subsequently increase N loss via gaseous and soil N-leaching. These findings bring us a step closer to predicting the responses and feedbacks of microbial-mediated N-cycling processes to long-term anthropogenic N-addition in grasslands.

Human activities are enriching many of Earth's ecosystems with biologically limiting mineral nutrients such as nitrogen (N) and phosphorus (P). In grasslands, this enrichment generally reduces plant diversity and increases productivity. The widely demonstrated positive effect of diversity on productivity suggests a potential negative feedback, whereby nutrient-induced declines in diversity reduce the initial gains in productivity arising from nutrient enrichment. In addition, plant productivity and diversity can be inhibited by accumulations of dead biomass, which may be altered by nutrient enrichment. Over longer time frames, nutrient addition may increase soil fertility by increasing soil organic matter and nutrient pools. We examined the effects of 5-11 yr of nutrient addition at 47 grasslands in 12 countries. Nutrient enrichment increased aboveground live biomass and reduced plant diversity at nearly all sites, and these effects became stronger over time. We did not find evidence that nutrient-induced losses of diversity reduced the positive effects of nutrients on biomass; however, nutrient effects on live biomass increased more slowly at sites where litter was also increasing, regardless of plant diversity. This work suggests that short-term experiments may underestimate the long-term nutrient enrichment effects on global grassland ecosystems.

Anthropogenic activities are increasing nutrient inputs to ecosystems worldwide, with consequences for global carbon and nutrient cycles. Recent meta-analyses show that aboveground primary production is often co-limited by multiple nutrients; however, little is known about how root production responds to changes in nutrient availability. At twenty-nine grassland sites on four continents, we quantified shallow root biomass responses to nitrogen (N), phosphorus (P) and potassium plus micronutrient enrichment and compared below- and aboveground responses. We hypothesized that optimal allocation theory would predict context dependence in root biomass responses to nutrient enrichment, given variation among sites in the resources limiting to plant growth (specifically light versus nutrients). Consistent with the predictions of optimal allocation theory, the proportion of total biomass belowground declined with N or P addition, due to increased biomass aboveground (for N and P) and decreased biomass belowground (N, particularly in sites with low canopy light penetration). Absolute root biomass increased with N addition where light was abundant at the soil surface, but declined in sites where the grassland canopy intercepted a large proportion of incoming light. These results demonstrate that belowground responses to changes in resource supply can differ strongly from aboveground responses, which could significantly modify predictions of future rates of nutrient cycling and carbon sequestration. Our results also highlight how optimal allocation theory developed for individual plants may help predict belowground biomass responses to nutrient enrichment at the ecosystem scale across wide climatic and environmental gradients.

Leaf traits are frequently measured in ecology to provide a ‘common currency’ for predicting how anthropogenic pressures impact ecosystem function. Here, we test whether leaf traits consistently respond to experimental treatments across 27 globally distributed grassland sites across 4 continents. We find that specific leaf area (leaf area per unit mass)—a commonly measured morphological trait inferring shifts between plant growth strategies—did not respond to up to four years of soil nutrient additions. Leaf nitrogen, phosphorus and potassium concentrations increased in response to the addition of each respective soil nutrient. We found few significant changes in leaf traits when vertebrate herbivores were excluded in the short-term. Leaf nitrogen and potassium concentrations were positively correlated with species turnover, suggesting that interspecific trait variation was a significant predictor of leaf nitrogen and potassium, but not of leaf phosphorus concentration. Climatic conditions and pretreatment soil nutrient levels also accounted for significant amounts of variation in the leaf traits measured. Overall, we find that leaf morphological traits, such as specific leaf area, are not appropriate indicators of plant response to anthropogenic perturbations in grasslands. Fil: Firn, Jennifer. Queensland University of Technology; Australia Fil: McGree, James M.. Queensland University of Technology; Australia Fil: Harvey, Eric. University of Montreal; Canadá Fil: Flores Moreno, Habacuc. University of Minnesota; Estados Unidos Fil: Schütz, Martin. Swiss Federal Institute for Forest, Snow and Landscape Research; Suiza Fil: Buckley, Yvonne M.. Trinity College; Irlanda Fil: Borer, Elizabeth T.. University of Minnesota; Estados Unidos Fil: Seabloom, Eric W.. University of Minnesota; Estados Unidos Fil: La Pierre, Kimberly J.. Smithsonian Environmental Research Center; Estados Unidos Fil: MacDougall, Andrew M.. University of Guelph; Canadá Fil: Prober, Suzanne M.. No especifíca; Fil: Stevens, Carly J.. Universidad de Lancaster; Reino Unido Fil: Sullivan, Lauren L.. University of Minnesota; Estados Unidos Fil: Porter, Erica. Queensland University of Technology; Australia Fil: Ladouceur, Emma. Helmholtz Center for Environmental Research; Alemania Fil: Allen, Charlotte. Queensland University of Technology; Australia Fil: Moromizato, Karine H.. Queensland University of Technology; Australia Fil: Morgan, John W.. La Trobe University; Australia Fil: Harpole, W. Stanley. Helmholtz Center for Environmental Research; Alemania. Universidad de Halle-Wittenberg ; Alemania Fil: Hautier, Yann. University of Utrecht; Países Bajos Fil: Eisenhauer, Nico. Universitat Leipzig; Alemania Fil: Wright, Justin P.. University of Duke; Estados Unidos Fil: Adler, Peter B.. State University of Utah; Estados Unidos Fil: Arnillas, Carlos Alberto. University of Toronto; Canadá Fil: Bakker, Jonathan D.. University of Washington; Estados Unidos Fil: Biederman, Lori. University of Iowa; Estados Unidos Fil: Broadbent, Arthur A. D.. University of Manchester; Reino Unido Fil: Brown, Cynthia S.. State University of Colorado at Boulder; Estados Unidos Fil: Bugalho, Miguel N.. Universidade Nova de Lisboa; Portugal Fil: Caldeira, Maria C.. Universidade Nova de Lisboa; Portugal Fil: Cleland, Elsa E.. University of California; Estados Unidos Fil: Ebeling, Anne. Universitat Jena; Alemania Fil: Fay, Philip A.. No especifíca; Fil: Hagenah, Nicole. Universidad de Pretoria; Sudáfrica Fil: Kleinhesselink, Andrew R.. University of California at Los Angeles; Estados Unidos Fil: Mitchell, Rachel. Arizona State University; Estados Unidos Fil: Moore, Joslin L.. Monash University; Australia Fil: Nogueira, Carla. Universidade Nova de Lisboa; Portugal Fil: Peri, Pablo Luis. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro de Investigaciones y Transferencia de Santa Cruz. Universidad Tecnológica Nacional. Facultad Regional Santa Cruz. Centro de Investigaciones y Transferencia de Santa Cruz. Universidad Nacional de la Patagonia Austral. Centro de Investigaciones y Transferencia de Santa Cruz; Argentina Fil: Roscher, Christiane. Helmholtz Center for Environmental Research; Alemania Fil: Smith, Melinda D.. State University of Colorado at Boulder; Estados Unidos Fil: Wragg, Peter D.. University of Minnesota; Estados Unidos Fil: Risch, Anita C.. No especifíca

Grasslands are subject to considerable alteration due to human activities globally, including widespread changes in populations and composition of large mammalian herbivores and elevated supply of nutrients. Grassland soils remain important reservoirs of carbon (C) and nitrogen (N). Herbivores may affect both C and N pools and these changes likely interact with increases in soil nutrient availability. Given the scale of grassland soil fluxes, such changes can have striking consequences for atmospheric C concentrations and the climate. Here, we use the Nutrient Network experiment to examine the responses of soil C and N pools to mammalian herbivore exclusion across 22 grasslands, under ambient and elevated nutrient availabilities (fertilized with NPK + micronutrients). We show that the impact of herbivore exclusion on soil C and N pools depends on fertilization. Under ambient nutrient conditions, we observed no effect of herbivore exclusion, but under elevated nutrient supply, pools are smaller upon herbivore exclusion. The highest mean soil C and N pools were found in grazed and fertilized plots. The decrease in soil C and N upon herbivore exclusion in combination with fertilization correlated with a decrease in aboveground plant biomass and microbial activity, indicating a reduced storage of organic matter and microbial residues as soil C and N. The response of soil C and N pools to herbivore exclusion was contingent on temperature – herbivores likely cause losses of C and N in colder sites and increases in warmer sites. Additionally, grasslands that contain mammalian herbivores have the potential to sequester more N under increased temperature variability and nutrient enrichment than ungrazed grasslands. Our study highlights the importance of conserving mammalian herbivore populations in grasslands worldwide. We need to incorporate local‐scale herbivory, and its interaction with nutrient enrichment and climate, within global‐scale models to better predict land–atmosphere interactions under future climate change., Global Change Biology, 26 (4), ISSN:1354-1013, ISSN:1365-2486

Fraser et al. (Reports, 17 July 2015, p. 302) report a unimodal relationship between productivity and species richness at regional and global scales, which they contrast with the results of Adler et al. (Reports, 23 September 2011, p. 1750). However, both data sets, when analyzed correctly, show clearly and consistently that productivity is a poor predictor of local species richness.