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1. Sensitivity of grassland carbon pools to plant diversity, elevated CO ₂ , and soil nitrogen addition over 19 years

   https://doi.org/10.1073/pnas.2016965118  

   Pastore, Melissa A.; Hobbie, Sarah E.; Reich, Peter B. (April 2021, Proceedings of the National Academy of Sciences)

   Whether the terrestrial biosphere will continue to act as a net carbon (C) sink in the face of multiple global changes is questionable. A key uncertainty is whether increases in plant C fixation under elevated carbon dioxide (CO₂) will translate into decades-long C storage and whether this depends on other concurrently changing factors. We investigated how manipulations of CO₂, soil nitrogen (N) supply, and plant species richness influenced total ecosystem (plant + soil to 60 cm) C storage over 19 y in a free-air CO₂enrichment grassland experiment (BioCON) in Minnesota. On average, after 19 y of treatments, increasing species richness from 1 to 4, 9, or 16 enhanced total ecosystem C storage by 22 to 32%, whereas N addition of 4 g N m⁻²⋅ y⁻¹and elevated CO₂of +180 ppm had only modest effects (increasing C stores by less than 5%). While all treatments increased net primary productivity, only increasing species richness enhanced net primary productivity sufficiently to more than offset enhanced C losses and substantially increase ecosystem C pools. Effects of the three global change treatments were generally additive, and we did not observe any interactions between CO₂and N. Overall, our results call into question whether elevated CO₂will increase the soil C sink in grassland ecosystems, helping to slow climate change, and suggest that losses of biodiversity may influence C storage as much as or more than increasing CO₂or high rates of N deposition in perennial grassland systems. 

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2. Role of external inputs of nutrients to aquatic ecosystems in determining prevalence of nitrogen vs. phosphorus limitation of net primary productivity

   https://doi.org/10.1007/s10533-021-00765-z  

   Howarth, R. W.; Chan, F.; Swaney, D. P.; Marino, R. M.; Hayn, M. (February 2021, Biogeochemistry)

   null (Ed.)

   Abstract Whether net primary productivity in an aquatic ecosystem is limited by nitrogen (N), limited by phosphorus (P), or co-limited by N & P is determined by the relative supply of N and P to phytoplankton compared to their elemental requirements for primary production, often characterized by the “Redfield” ratio. The supply of these essential nutrients is affected by both external inputs and biogeochemical processes within the ecosystem. In this paper, we examine external sources of nutrients to aquatic systems and how the balance of N to P inputs influences nutrient limitation. For ocean subtropical gyres, a relatively balanced input of N and P relative to the Redfield ratio from deep ocean sources often leads to near co-limitation by N and P. For lakes, the external nutrient inputs come largely from watershed sources, and we demonstrate that on average the N:P ratio for these inputs across the United States is well above that needed by phytoplankton, which may contribute to P limitation in those lake that experience this average nutrient loading. Watershed inputs are also important for estuaries and coastal marine ecosystems, but ocean sources of nutrients are also significant contributors to overall nutrient loads. The ocean-nutrient sources of N and P are very often at or below the Redfield ratio of 16:1 molar, and can be substantially so, particularly in areas where the continental shelf is wide. This large input of coastal ocean nutrients with a low N:P ratio is one factor that may make N limitation more likely in many coastal marine ecosystems than in lakes. 

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3. Evidence, causes, and consequences of declining nitrogen availability in terrestrial ecosystems

   https://doi.org/10.1126/science.abh3767  

   Mason, Rachel E.; Craine, Joseph M.; Lany, Nina K.; Jonard, Mathieu; Ollinger, Scott V.; Groffman, Peter M.; Fulweiler, Robinson W.; Angerer, Jay; Read, Quentin D.; Reich, Peter B.; et al (April 2022, Science)

   BACKGROUND The availability of nitrogen (N) to plants and microbes has a major influence on the structure and function of ecosystems. Because N is an essential component of plant proteins, low N availability constrains the growth of plants and herbivores. To increase N availability, humans apply large amounts of fertilizer to agricultural systems. Losses from these systems, combined with atmospheric deposition of fossil fuel combustion products, introduce copious quantities of reactive N into ecosystems. The negative consequences of these anthropogenic N inputs—such as ecosystem eutrophication and reductions in terrestrial and aquatic biodiversity—are well documented. Yet although N availability is increasing in many locations, reactive N inputs are not evenly distributed globally. Furthermore, experiments and theory also suggest that global change factors such as elevated atmospheric CO 2 , rising temperatures, and altered precipitation and disturbance regimes can reduce the availability of N to plants and microbes in many terrestrial ecosystems. This can occur through increases in biotic demand for N or reductions in its supply to organisms. Reductions in N availability can be observed via several metrics, including lowered nitrogen concentrations ([N]) and isotope ratios (δ 15 N) in plant tissue, reduced rates of N mineralization, and reduced terrestrial N export to aquatic systems. However, a comprehensive synthesis of N availability metrics, outside of experimental settings and capable of revealing large-scale trends, has not yet been carried out. ADVANCES A growing body of observations confirms that N availability is declining in many nonagricultural ecosystems worldwide. Studies have demonstrated declining wood δ 15 N in forests across the continental US, declining foliar [N] in European forests, declining foliar [N] and δ 15 N in North American grasslands, and declining [N] in pollen from the US and southern Canada. This evidence is consistent with observed global-scale declines in foliar δ 15 N and [N] since 1980. Long-term monitoring of soil-based N availability indicators in unmanipulated systems is rare. However, forest studies in the northeast US have demonstrated decades-long decreases in soil N cycling and N exports to air and water, even in the face of elevated atmospheric N deposition. Collectively, these studies suggest a sustained decline in N availability across a range of terrestrial ecosystems, dating at least as far back as the early 20th century. Elevated atmospheric CO 2 levels are likely a main driver of declines in N availability. Terrestrial plants are now uniformly exposed to ~50% more of this essential resource than they were just 150 years ago, and experimentally exposing plants to elevated CO 2 often reduces foliar [N] as well as plant-available soil N. In addition, globally-rising temperatures may raise soil N supply in some systems but may also increase N losses and lead to lower foliar [N]. Changes in other ecosystem drivers—such as local climate patterns, N deposition rates, and disturbance regimes—individually affect smaller areas but may have important cumulative effects on global N availability. OUTLOOK Given the importance of N to ecosystem functioning, a decline in available N is likely to have far-reaching consequences. Reduced N availability likely constrains the response of plants to elevated CO 2 and the ability of ecosystems to sequester carbon. Because herbivore growth and reproduction scale with protein intake, declining foliar [N] may be contributing to widely reported declines in insect populations and may be negatively affecting the growth of grazing livestock and herbivorous wild mammals. Spatial and temporal patterns in N availability are not yet fully understood, particularly outside of Europe and North America. Developments in remote sensing, accompanied by additional historical reconstructions of N availability from tree rings, herbarium specimens, and sediments, will show how N availability trajectories vary among ecosystems. Such assessment and monitoring efforts need to be complemented by further experimental and theoretical investigations into the causes of declining N availability, its implications for global carbon sequestration, and how its effects propagate through food webs. Responses will need to involve reducing N demand via lowering atmospheric CO 2 concentrations, and/or increasing N supply. Successfully mitigating and adapting to declining N availability will require a broader understanding that this phenomenon is occurring alongside the more widely recognized issue of anthropogenic eutrophication. Intercalibration of isotopic records from leaves, tree rings, and lake sediments suggests that N availability in many terrestrial ecosystems has steadily declined since the beginning of the industrial era. Reductions in N availability may affect many aspects of ecosystem functioning, including carbon sequestration and herbivore nutrition. Shaded areas indicate 80% prediction intervals; marker size is proportional to the number of measurements in each annual mean. Isotope data: (tree ring) K. K. McLauchlan et al. , Sci. Rep. 7 , 7856 (2017); (lake sediment) G. W. Holtgrieve et al. , Science 334 , 1545–1548 (2011); (foliar) J. M. Craine et al. , Nat. Ecol. Evol. 2 , 1735–1744 (2018) 

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4. Acclimation of Photosynthesis to CO ₂ Increases Ecosystem Carbon Storage due to Leaf Nitrogen Savings

   https://doi.org/10.1111/gcb.17558  

   Smith, Nicholas G; Zhu, Qing; Keenan, Trevor F; Riley, William J (November 2024, Global Change Biology)

   ABSTRACT Photosynthesis is the largest flux of carbon between the atmosphere and Earth's surface and is driven by enzymes that require nitrogen, namely, ribulose‐1,5‐bisphosphate (RuBisCO). Thus, photosynthesis is a key link between the terrestrial carbon and nitrogen cycle, and the representation of this link is critical for coupled carbon‐nitrogen land surface models. Models and observations suggest that soil nitrogen availability can limit plant productivity increases under elevated CO₂. Plants acclimate to elevated CO₂by downregulating RuBisCO and thus nitrogen in leaves, but this acclimation response is not currently included in land surface models. Acclimation of photosynthesis to CO₂can be simulated by the photosynthetic optimality theory in a way that matches observations. Here, we incorporated this theory into the land surface component of the Energy Exascale Earth System Model (ELM). We simulated land surface carbon and nitrogen processes under future elevated CO₂conditions to 2100 using the RCP8.5 high emission scenario. Our simulations showed that when photosynthetic acclimation is considered, photosynthesis increases under future conditions, but maximum RuBisCO carboxylation and thus photosynthetic nitrogen demand decline. We analyzed two simulations that differed as to whether the saved nitrogen could be used in other parts of the plant. The allocation of saved leaf nitrogen to other parts of the plant led to (1) a direct alleviation of plant nitrogen limitation through reduced leaf nitrogen requirements and (2) an indirect reduction in plant nitrogen limitation through an enhancement of root growth that led to increased plant nitrogen uptake. As a result, reallocation of saved leaf nitrogen increased ecosystem carbon stocks by 50.3% in 2100 as compared to a simulation without reallocation of saved leaf nitrogen. These results suggest that land surface models may overestimate future ecosystem nitrogen limitation if they do not incorporate leaf nitrogen savings resulting from photosynthetic acclimation to elevated CO₂. 

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5. Flexible Foliar Stoichiometry Reduces the Magnitude of the Global Land Carbon Sink

   https://doi.org/10.1029/2023GL105493  

   Hauser, Emma; Wieder, William_R; Bonan, Gordon_B; Cleveland, Cory_C (November 2023, Geophysical Research Letters)

   Abstract Increased plant growth under elevated carbon dioxide (CO₂) slows the pace of climate warming and underlies projections of terrestrial carbon (C) and climate dynamics. However, this important ecosystem service may be diminished by concurrent changes to vegetation carbon‐to‐nitrogen (C:N) ratios. Despite clear observational evidence of increasing foliar C:N under elevated CO₂, our understanding of potential ecological consequences of foliar stoichiometric flexibility is incomplete. Here, we illustrate that when we incorporated CO₂‐driven increases in foliar stoichiometry into the Community Land Model the projected land C sink decreased two‐fold by the end of the century compared to simulations with fixed foliar chemistry. Further, CO₂‐driven increases in foliar C:N profoundly altered Earth's hydrologic cycle, reducing evapotranspiration and increasing runoff, and reduced belowground N cycling rates. These findings underscore the urgency of further research to examine both the direct and indirect effects of changing foliar stoichiometry on soil N cycling and plant productivity. 

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