Patterns and controls of foliar nutrient stoichiometry and flexibility across United States forests.

Plant element stoichiometry and stoichiometric flexibility strongly regulate ecosystem responses to global change. Here, we tested three potential mechanistic drivers (climate, soil nutrients, and plant taxonomy) of both using paired foliar and soil nutrient data from terrestrial forested National Ecological Observatory Network sites across the USA. We found that broad patterns of foliar nitrogen (N) and foliar phosphorus (P) are explained by different mechanisms. Plant taxonomy was an important control over all foliar nutrient stoichiometries and concentrations, especially foliar N, which was dominantly related to taxonomy and did not vary across climate or soil gradients. Despite a lack of site-level correlations between N and environment variables, foliar N exhibited intraspecific flexibility, with numerous species-specific correlations between foliar N and various environmental factors, demonstrating the variable spatial and temporal scales on which foliar chemistry and stoichiometric flexibility can manifest. In addition to plant taxonomy, foliar P and N:P ratios were also linked to soil nutrient status (extractable P) and climate, especially actual evapotranspiration rates. Our findings highlight the myriad factors that influence foliar chemistry and show that broad patterns cannot be explained by a single consistent mechanism. Furthermore, differing controls over foliar N versus P suggests that each may be sensitive to global change drivers on distinct spatial and temporal scales, potentially resulting in altered ecosystem N:P ratios that have implications for processes ranging from productivity to carbon sequestration.

Isotopic constraints on plant nitrogen acquisition strategies during ecosystem retrogression.

Plant root associations with microbes such as mycorrhizal fungi or N-fixing bacteria enable ecosystems to tap pools of nitrogen (N) that might otherwise be inaccessible, including atmospheric N or N in large soil organic molecules. Such microbially assisted N-foraging strategies may be particularly important in late-successional retrogressive ecosystems where productivity is low and soil nutrients are scarce. Here, we use natural N-stable isotopic composition to constrain pathways of N supplies to different plant functional groups across a well-studied natural soil fertility gradient that includes a highly retrogressive stage. We demonstrate that ectomycorrhizal fungi, ericoid mycorrhizal fungi, and N-fixing bacteria support forest N supplies at all stages of ecosystem succession, from relatively young, N-rich/phosphorus (P)-rich sites, to ancient sites (ca. 500 ky) where both N supplies and P supplies are exceedingly low. Microbially mediated N sources are most important in older ecosystems with very low soil nutrient availability, accounting for 75-96% of foliar N at the oldest, least fertile sites. These isotopically ground findings point to the key role of plant-microbe associations in shaping ecosystem processes and functioning, particularly in retrogressive-phase forest ecosystems.

Bedrock nitrogen weathering stimulates biological nitrogen fixation.

Global ecosystem models suggest that bedrock nitrogen (N) weathering contributes 10-20% of total N inputs to the natural terrestrial biosphere and >38% of ecosystem N supplies in temperate forests specifically. Yet, the role of rock N weathering in shaping ecological processes and biogeochemical fluxes is largely unknown. Here, we show that temperate forest ecosystems underlain by N-rich bedrock exhibit higher free-living N fixation rates than similar forests residing on N-poor parent materials, across sites experiencing a range of climate and tectonic regimes. This seemingly counterintuitive result can be explained by increased accumulation of soil C and P in high bedrock N sites, resulting in increased energy inputs and nutrient supplies to N fixing microorganisms. Our findings advance a novel ecosystem biogeochemical framework that recognizes long-term plant-soil-microbe feedbacks in shaping biogeochemical processes, with potentially widespread implications given the global distribution of bedrock N across Earth's terrestrial biomes.

Nutrient limitation of terrestrial free-living nitrogen fixation.

Nitrogen (N) fixation by free-living bacteria is a primary N input pathway in many ecosystems and sustains global plant productivity. Uncertainty exists over the importance of N, phosphorus (P) and molybdenum (Mo) availability in controlling free-living N fixation rates. Here, we investigate the geographic occurrence and variability of nutrient constraints to free-living N fixation in the terrestrial biosphere. We compiled data from studies measuring free-living N fixation in response to N, P and Mo fertilizers. We used meta-analysis to quantitatively determine the extent to which N, P and Mo stimulate or suppress N fixation, and if environmental variables influence the degree of nutrient limitation of N fixation. Across our compiled dataset, free-living N fixation is suppressed by N fertilization and stimulated by Mo fertilization. Additionally, free-living N fixation is stimulated by P additions in tropical forests. These findings suggest that nutrient limitation is an intrinsic property of the biochemical demands of N fixation, constraining free-living N fixation in the terrestrial biosphere. These findings have implications for understanding the causes and consequences of N limitation in coupled nutrient cycles, as well as modeling and forecasting nutrient controls over carbon-climate feedbacks.