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1. Negative effects of nitrogen override positive effects of phosphorus on grassland legumes worldwide

   https://doi.org/10.1073/pnas.2023718118  

   Tognetti, Pedro M.; Prober, Suzanne M.; Báez, Selene; Chaneton, Enrique J.; Firn, Jennifer; Risch, Anita C.; Schuetz, Martin; Simonsen, Anna K.; Yahdjian, Laura; Borer, Elizabeth T.; et al (July 2021, Proceedings of the National Academy of Sciences)

   null (Ed.)

   Anthropogenic nutrient enrichment is driving global biodiversity decline and modifying ecosystem functions. Theory suggests that plant functional types that fix atmospheric nitrogen have a competitive advantage in nitrogen-poor soils, but lose this advantage with increasing nitrogen supply. By contrast, the addition of phosphorus, potassium, and other nutrients may benefit such species in low-nutrient environments by enhancing their nitrogen-fixing capacity. We present a global-scale experiment confirming these predictions for nitrogen-fixing legumes (Fabaceae) across 45 grasslands on six continents. Nitrogen addition reduced legume cover, richness, and biomass, particularly in nitrogen-poor soils, while cover of non–nitrogen-fixing plants increased. The addition of phosphorous, potassium, and other nutrients enhanced legume abundance, but did not mitigate the negative effects of nitrogen addition. Increasing nitrogen supply thus has the potential to decrease the diversity and abundance of grassland legumes worldwide regardless of the availability of other nutrients, with consequences for biodiversity, food webs, ecosystem resilience, and genetic improvement of protein-rich agricultural plant species. 

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2. Diversity‐dependent soil acidification under nitrogen enrichment constrains biomass productivity

   https://doi.org/10.1111/gcb.15329  

   Kimmel, Kaitlin; Furey, George N.; Hobbie, Sarah E.; Isbell, Forest; Tilman, David; Reich, Peter B. (September 2020, Global Change Biology)

   Abstract In most plant communities, the net effect of nitrogen enrichment is an increase in plant productivity. However, nitrogen enrichment also has been shown to decrease species richness and to acidify soils, each of which may diminish the long‐term impact of nutrient enrichment on productivity. Here we use a long‐term (20 year) grassland plant diversity by nitrogen enrichment experiment in Minnesota, United States (a subexperiment within the BioCON experiment) to quantify the net impacts of nitrogen enrichment on productivity, including its potential indirect effects on productivity via changes in species richness and soil pH over an experimental diversity gradient. Overall, we found that nitrogen enrichment led to an immediate positive increment in productivity, but that this effect became nonsignificant over later years of the experiment, with the difference in productivity between fertilized and unfertilized plots decreasing in proportion to nitrogen addition‐dependent declines in soil pH and losses of plant diversity. The net effect of nitrogen enrichment on productivity could have been 14.5% more on average over 20 years in monocultures if not for nitrogen‐induced decreases in pH and about 28.5% more on average over 20 years in 16 species communities if not for nitrogen‐induced species richness losses. Together, these results suggest that the positive effects of nutrient enrichment on biomass production can diminish in their magnitude over time, especially because of soil acidification in low diversity communities and especially because of plant diversity loss in initially high diversity communities. 

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3. Interactions among nutrients govern the global grassland biomass–precipitation relationship

   https://doi.org/10.1073/pnas.2410748122  

   Fay, Philip A; Gherardi, Laureano A; Yahdjian, Laura; Adler, Peter B; Bakker, Jonathan D; Bharath, Siddharth; Borer, Elizabeth T; Harpole, W Stanley; Hersch-Green, Erika; Huxman, Travis E; et al (April 2025, Proceedings of the National Academy of Sciences)

   Ecosystems are experiencing changing global patterns of mean annual precipitation (MAP) and enrichment with multiple nutrients that potentially colimit plant biomass production. In grasslands, mean aboveground plant biomass is closely related to MAP, but how this relationship changes after enrichment with multiple nutrients remains unclear. We hypothesized the global biomass–MAP relationship becomes steeper with an increasing number of added nutrients, with increases in steepness corresponding to the form of interaction among added nutrients and with increased mediation by changes in plant community diversity. We measured aboveground plant biomass production and species diversity in 71 grasslands on six continents representing the global span of grassland MAP, diversity, management, and soils. We fertilized all sites with nitrogen, phosphorus, and potassium with micronutrients in all combinations to identify which nutrients limited biomass at each site. As hypothesized, fertilizing with one, two, or three nutrients progressively steepened the global biomass–MAP relationship. The magnitude of the increase in steepness corresponded to whether sites were not limited by nitrogen or phosphorus, were limited by either one, or were colimited by both in additive, or synergistic forms. Unexpectedly, we found only weak evidence for mediation of biomass–MAP relationships by plant community diversity because relationships of species richness, evenness, and beta diversity to MAP and to biomass were weak or opposing. Site-level properties including baseline biomass production, soils, and management explained little variation in biomass–MAP relationships. These findings reveal multiple nutrient colimitation as a defining feature of the global grassland biomass–MAP relationship. 

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4. Nutrient identity modifies the destabilising effects of eutrophication in grasslands

   https://doi.org/10.1111/ele.13946  

   Carroll, Oliver; Batzer, Evan; Bharath, Siddharth; Borer, Elizabeth T.; Campana, Sofía; Esch, Ellen; Hautier, Yann; Ohlert, Timothy; Seabloom, Eric W.; Adler, Peter B.; et al (December 2021, Ecology Letters)

   Abstract Nutrient enrichment can simultaneously increase and destabilise plant biomass production, with co‐limitation by multiple nutrients potentially intensifying these effects. Here, we test how factorial additions of nitrogen (N), phosphorus (P) and potassium with essential nutrients (K+) affect the stability (mean/standard deviation) of aboveground biomass in 34 grasslands over 7 years. Destabilisation with fertilisation was prevalent but was driven by single nutrients, not synergistic nutrient interactions. On average, N‐based treatments increased mean biomass production by 21–51% but increased its standard deviation by 40–68% and so consistently reduced stability. Adding P increased interannual variability and reduced stability without altering mean biomass, while K+ had no general effects. Declines in stability were largest in the most nutrient‐limited grasslands, or where nutrients reduced species richness or intensified species synchrony. We show that nutrients can differentially impact the stability of biomass production, with N and P in particular disproportionately increasing its interannual variability. 

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5. Plant diversity and litter accumulation mediate the loss of foliar endophyte fungal richness following nutrient addition

   https://doi.org/10.1002/ecy.3210  

   Henning, Jeremiah A.; Kinkel, Linda; May, Georgiana; Lumibao, Candice Y.; Seabloom, Eric W.; Borer, Elizabeth T. (December 2020, Ecology)

   Abstract Foliar fungal endophytes are ubiquitous plant symbionts that can affect plant growth and reproduction via their roles in pathogen and stress tolerance, as well as plant hormonal signaling. Despite their importance, we have a limited understanding of how foliar fungal endophytes respond to varying environmental conditions such as nutrient inputs. The responses of foliar fungal endophyte communities to increased nutrient deposition may be mediated by the simultaneous effects on within‐host competition as well as the indirect impacts of altered host population size, plant productivity, and plant community diversity and composition. Here, we leveraged a 7‐yr experiment manipulating nitrogen, phosphorus, potassium, and micronutrients to investigate how nutrient‐induced changes to plant diversity, plant productivity, and plant community composition relate to changes in foliar fungal endophyte diversity and richness in a focal native grass host,Andropogon gerardii. We found limited evidence of direct effects of nutrients on endophyte diversity. Instead, the effects of nutrients on endophyte diversity appeared to be mediated by accumulation of plant litter and plant diversity loss. Specifically, nitrogen addition is associated with a 40% decrease in plant diversity and an 11% decrease in endophyte richness. Although nitrogen, phosphorus, and potassium addition increased aboveground live biomass and decreased relativeAndropogoncover, endophyte diversity did not covary with live plant biomass orAndropogoncover. Our results suggest that fungal endophyte diversity within this focal host is determined in part by the diversity of the surrounding plant community and its potential impact on immigrant propagules and dispersal dynamics. Our results suggest that elemental nutrients reduce endophyte diversity indirectly via impacts on the local plant community, not direct response to nutrient addition. Thus, the effects of global change drivers, such as nutrient deposition, on characteristics of host populations and the diversity of their local communities are important for predicting the response of symbiont communities in a changing global environment. 

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