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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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Nitrogen Deposition and Terrestrial Biodiversity
Nitrogen deposition, along with habitat losses and climate change, has been identified as a primary threat to biodiversity worldwide (Butchart et al., 2010; MEA, 2005; Sala et al., 2000). The source of this stressor to natural systems is generally twofold: burning of fossil fuels and the use of fertilizers in modern intensive agriculture. Each of these human enterprises leads to the release of large amounts of biologically reactive nitrogen (henceforth contracted to "nitrogen") to the atmosphere, which is later deposited to ecosystems. Because nitrogen is a critical element to all living things (as a primary building block of proteins among other biological molecules), nitrogen availability often limits primary production and is tightly recycled in many natural ecosystems. This is especially true in temperate ecosystems, though it may also be true for some areas in the tropics that are not phosphorus-limited (Adams et al., 2004; Matson et al., 1999). Thus, the large increase in availability of this critical nutrient as a result of human activity has profound impacts on ecosystems and on biodiversity. Once nitrogen is deposited on terrestrial ecosystems, a cascade of effects can occur that often leads to overall declines in biodiversity (Bobbink et al., 2010; Galloway et al., 2003). For plants, nitrogen deposition can impact biodiversity generally through four processes: (1) stimulation of growth often of weedy species that outcompete local neighbors (termed "eutrophication"), (2) acidification of the soil and consequent imbalances in other key nutrients that favors acid tolerant species (termed "acidification"), (3) enhancement of secondary stressors such as from fire, drought, frost, or pests triggered by increased nitrogen availability (termed "secondary stressors"), and (4) direct damage to leaves (termed "direct toxicity") (Bobbink, 1998; Bobbink et al., 2010). For animals, much less is known, but reductions in plant biodiversity can lead to reductions in diversity of invertebrate and other animal species, loss of habitat heterogeneity and specialist habitats, increased pest populations and activity, and changes in soil microbial communities (McKinney and Lockwood, 1999; Throop and Lerdau, 2004; Treseder, 2004). Citation Clark, Christopher M.; Bai, Yongfei; Bowman, William D.; Cowles, Jane M.; Fenn, Mark E.; Gilliam, Frank S.; Phoenix, Gareth K.; Siddique, Ilyas; Stevens, Carly J.; Sverdrup, Harald U.; Throop, Heather L. 2013. Nitrogen deposition and terrestrial biodiversity. In: Levin S.A. (ed.) Encyclopedia of Biodiversity, second edition, Volume 5, Waltham, MA: Academic Press. pp. 519-536.
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- Award ID(s):
- 2425290
- PAR ID:
- 10654322
- Publisher / Repository:
- Elsevier
- Date Published:
- ISSN:
- 978‑0123847195
- Page Range / eLocation ID:
- 519 to 536
- Format(s):
- Medium: X
- Sponsoring Org:
- National Science Foundation
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- https://doi.org/10.1016/B978-0-12-384719-5.00366-X
