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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. Phosphorus deprivation affects composition and spatial distribution of membrane lipids in legume nodules.

   https://doi.org/doi.org/10.1093/plphys/kiaa115  

   Dokwal, D; Romsdahl, TB; Kunz, DA; Alonso, AP; Dickstein, R (January 2021, Plant physiology)

   null (Ed.)

   In legumes, symbiotic nitrogen (N) fixation (SNF) occurs in specialized organs called nodules after successful interactions between legume hosts and rhizobia. In a nodule, N-fixing rhizobia are surrounded by symbiosome membranes, through which the exchange of nutrients and ammonium occurs between bacteria and the host legume. Phosphorus (P) is an essential macronutrient, and N2-fixing legumes have a higher requirement for P than legumes grown on mineral N. As in the previous studies, in P deficiency, barrel medic (Medicago truncatula) plants had impaired SNF activity, reduced growth, and accumulated less phosphate in leaves, roots, and nodules compared with the plants grown in P sufficient conditions. Membrane lipids in M. truncatula tissues were assessed using electrospray ionization–mass spectrometry. Galactolipids were found to increase in P deficiency, with declines in phospholipids (PL), especially in leaves. Lower PL losses were found in roots and nodules. Subsequently, matrix-assisted laser desorption/ionization–mass spectrometry imaging was used to spatially map the distribution of the positively charged phosphatidylcholine (PC) species in nodules in both P-replete and P-deficient conditions. Our results reveal heterogeneous distribution of several PC species in nodules, with homogeneous distribution of other PC classes. In P poor conditions, some PC species distributions were observed to change. The results suggest that specific PC species may be differentially important in diverse nodule zones and cell types, and that membrane lipid remodeling during P stress is not uniform across the nodule. 

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3. Legumes regulate grassland soil N cycling and its response to variation in species diversity and N supply but not CO ₂

   https://doi.org/10.1111/gcb.14636  

   Wei, Xiaorong; Reich, Peter_B; Hobbie, Sarah_E (April 2019, Global Change Biology)

   Abstract Legumes are an important component of plant diversity that modulate nitrogen (N) cycling in many terrestrial ecosystems. Limited knowledge of legume effects on soil N cycling and its response to global change factors and plant diversity hinders a general understanding of whether and how legumes broadly regulate the response of soil N availability to those factors. In a 17‐year study of perennial grassland species grown under ambient and elevated (+180 ppm) CO₂and ambient and enriched (+4 g N m⁻² year⁻¹) N environments, we compared pure legume plots with plots dominated by or including other herbaceous functional groups (and containing one or four species) to assess the effect of legumes on N cycling (net N mineralization rate and inorganic N pools). We also examined the effects of numbers of legume species (from zero to four) in four‐species mixed plots on soil N cycling. We hypothesized that legumes would increase N mineralization rates most in those treatments with the greatest diversity and the greatest relative limitation by and competition for N. Results partially supported these hypotheses. Plots with greater dominance by legumes had greater soil nitrate concentrations and mineralization rates. Higher species richness significantly increased the impact of legumes on soil N metrics, with 349% and 505% higher mineralization rates and nitrate concentrations in four‐species plots containing legumes compared to legume‐free four‐species plots, in contrast to 185% and 129% greater values, respectively, in pure legume than nonlegume monoculture plots. N‐fertilized plots had greater legume effects on soil nitrate, but lower legume effects on net N mineralization. In contrast, neither elevated CO₂nor its interaction with legumes affected net N mineralization. These results indicate that legumes markedly influence the response of soil N cycling to some, but not all, global change drivers. 

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4. Metabolite shift in Medicago truncatula occurs in phosphorus deprivation

   https://doi.org/10.1093/jxb/erab559  

   Dokwal, Dhiraj; Cocuron, Jean-Christophe; Alonso, Ana Paula; Dickstein, Rebecca; Nakamura, ed., Yuki (December 2021, Journal of Experimental Botany)

   Abstract Symbiotic nitrogen (N) fixation entails successful interaction between legume hosts and rhizobia that occur in specialized organs called nodules. N-fixing legumes have a higher demand for phosphorus (P) than legumes grown on mineral N. Medicago truncatula is an important model plant for characterization of effects of P deficiency at the molecular level. Hence, a study was carried out to address the alteration in metabolite levels of M. truncatula grown aeroponically and subjected to 4 weeks of P stress. First, GC-MS-based untargeted metabolomics initially revealed changes in the metabolic profile of nodules, with increased levels of amino acids and sugars and a decline in amounts of organic acids. Subsequently, LC-MS/MS was used to quantify these compounds including phosphorylated metabolites in the whole plant. Our results showed a drastic reduction in levels of organic acids and phosphorylated compounds in –P leaves, with a moderate reduction in –P roots and nodules. Additionally, sugars and amino acids were elevated in the whole plant under P deprivation. These findings provide evidence that N fixation in M. truncatula is mediated through a N feedback mechanism that in parallel is related to carbon and P metabolism. 

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5. Community change can buffer chronic nitrogen impacts, but multiple nutrients tip the scale

   https://doi.org/10.1002/ecy.3355  

   Wilcots, Megan E.; Harpole, W. Stanley; Seabloom, Eric W.; Borer, Elizabeth T. (May 2021, Ecology)

   Abstract Anthropogenic nitrogen (N) inputs are causing large changes in ecosystems worldwide. Many previous studies have examined the impact of N on terrestrial ecosystems; however, most have added N at rates that are much higher than predicted future deposition rates. Here, we present the results from a gradient of experimental N addition (0–10 g·N·m⁻²) in a temperate grassland. After a decade of N addition, we found that all levels of N addition changed plant functional group composition, likely indicating altered function for plant communities exposed to 10 yr of N inputs. However, N addition only had weak impacts on species composition and this functional group shift was not driven by any particular species, suggesting high levels of functional redundancy among grasslands species. Adding other nutrients (P, K, and micronutrients) in combination with N caused substantially greater changes in the relative abundance of species and functional groups. Together, these results suggest that compositional change within functional groups may buffer grasslands from impacts of N deposition, but concurrent eutrophication with other elements will likely lead to substantial changes in plant composition and biomass. 

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