The global nitrogen cycle: Past,present and future

Food and energy production converts N₂ to reactive N species that cascade through environmental reservoirs and in the process impact human and ecosystem health. This presentation will examine the impact of increased N mobilization on the global N cycle by contrasting N distribution in the late-19th century with those of the late-20th century. The presentation will give a general overview of regional differences and will conclude with a projection of the global N cycle for 2050.

     

The global nitrogen cycle: Past, present and future

Food and energy production converts N2 to reactive N species that cascade through environmental reservoirs and in the process impact human and ecosystem health. This presentation will examine the impact of increased N mobilization on the global N cycle by contrasting N distribution in the late-19th century with those of the late-20th century. The presentation will give a general overview of regional differences and will conclude with a projection of the global N cycle for 2050.     

The global nitrogen cycle: Past, present and future

Food and energy production converts N2 to reactive N species that cascade through environmental reservoirs and in the process impact human and ecosystem health. This presentation will examine the impact of increased N mobilization on the global N cycle by contrasting N distribution in the late-19th century with those of the late-20th century. The presentation will give a general overview of regional differences and will conclude with a projection of the global N cycle for 2050.     

The Jared Site: A Comanche Burial at Fort Sill,Oklahoma

Abstract

A late-19th century ravine burial from Southwest Oklahoma is described. The site is compared with historic and anthropological records to document a nineteenth century Comanche burial practice.     

Modelling terrestrial nitrous oxide emissions and implications for climate feedback

Ecosystem nitrous oxide (N(2) O) emissions respond to changes in climate and CO(2) concentration as well as anthropogenic nitrogen (N) enhancements. Here, we aimed to quantify the responses of natural ecosystem N(2) O emissions to multiple environmental drivers using a process-based global vegetation model (DyN-LPJ). We checked that modelled annual N(2) O emissions from nonagricultural ecosystems could reproduce field measurements worldwide, and experimentally observed responses to step changes in environmental factors. We then simulated global N(2) O emissions throughout the 20th century and analysed the effects of environmental changes. The model reproduced well the global pattern of N(2) O emissions and the observed responses of N cycle components to changes in environmental factors. Simulated 20th century global decadal-average soil emissions were c. 8.2-9.5?Tg?N?yr(-1) (or 8.3-10.3?Tg?N?yr(-1) with N deposition). Warming and N deposition contributed 0.85?±?0.41 and 0.80?±?0.14?Tg?N?yr(-1) , respectively, to an overall upward trend. Rising CO(2) also contributed, in part, through a positive interaction with warming. The modelled temperature dependence of N(2) O emission (c. 1?Tg?N?yr(-1) K(-1) ) implies a positive climate feedback which, over the lifetime of N(2) O (114?yr), could become as important as the climate-carbon cycle feedback caused by soil CO(2) release.     

A chronology of human understanding of the nitrogen cycle

Nitrogen over the ages! It was discovered in the eighteenth century. The following century, its importance in agriculture was documented and the basic components of its cycle were elucidated. In the twentieth century, a process to provide an inexhaustible supply of reactive N (N_r; all N species except N₂) for agricultural, industrial and military uses was invented. This discovery and the extensive burning of fossil fuels meant that by the beginning of the twenty-first century, anthropogenic sources of newly created N_r were two to three times that of natural terrestrial sources. This caused a fundamental change in the nitrogen cycle; for the first time, there was the potential for enough food to sustain growing populations and changing dietary patterns. However, most N_r created by humans is lost to the environment, resulting in a cascade of negative earth systems impacts—including enhanced acid rain, smog, eutrophication, greenhouse effect and stratospheric ozone depletion, with associated impacts on human and ecosystem health. The impacts continue and will be magnified, as N_r is lost to the environment at an even greater rate. Thus, the challenge for the current century is how to optimize the uses of N while minimizing the negative impacts.     

Exchange of gaseous nitrogen compounds between agricultural systems and the atmosphere

Crop and livestock agricultural production systems are important contributors to local, regional and global budgets of NH₃, NO_x (NO + NO₂) and N₂O. Emissions of NH₃ and NO_x (which are biologically and chemically active) into the atmosphere serve to redistribute fixed N to local and regional aquatic and terrestrial ecosystems that may otherwise be disconnected from the sources of the N gases. The emissions of NO_x also contribute to local elevated ozone concentrations while N₂O emissions contribute to global greenhouse gas accumulation and to stratospheric ozone depletion.Ammonia is the major gaseous base in the atmosphere and serves to neutralize about 30% of the hydrogen ions in the atmosphere. Fifty to 75% of the 55 Tg N yr^–1 NH₃ from terrestrial systems is emitted from animal and crop-based agriculture from animal excreta and synthetic fertilizer application. About half of the 50 Tg N yr^–1 of NO_x emitted from the earth surface annually arises from fossil fuel combustion and the remainder from biomass burning and emissions from soil. The NO_x emitted, principally as nitric oxide (NO), reacts rapidly in the atmosphere and in a complex cycle with light, ozone and hydrocarbons, and produces nitric acid and particulate nitrate. These materials can interact with plants and the soil locally or be transported form the site and interact with atmospheric particulate to form aerosols. These salts and aerosols return to fertilize terrestrial and aquatic systems in wet and dry deposition. A small fraction of this N may be biologically converted to N₂O. About 5% of the total atmospheric greenhouse effect is attributed to N₂O from which 70% of the annual global anthropogenic emissions come from animal and crop production.The coupling of increased population with a move of a large sector of the world population to diets that require more energy and N input, will lead to continued increases in anthropogenic input into the global N cycle. This scenario suggests that emissions of NH₃, NO_x and N₂O from agricultural systems will continue to increase and impact global terrestrial and aquatic systems, even those far removed from agricultural production, to an ever growing extent, unless N resources are used more efficiently or food consumption trends change.     

The biological standard of living in 19th century Mexico and in the American West

During the mid-19th century, the United States acquired Texas and large parts of Mexican territory with the vast Mexican-born population. This paper considers the biological standard of living of the part of this population that was incarcerated in American prisons. We use their physical stature as a proxy for their biological welfare. These data confirm earlier results which showed that adult heights tended to stagnate in Mexico during the late-19th century despite considerable social and political turmoil. While there is some evidence of a decline in height among youth, the decline is slight (<1 cm). As in other 19th century samples, farmers were the tallest. Americans were taller than Mexican prisoners by about 2 cm.     

土壤微生物对气候变暖和大气N沉降的响应

气候变暖和大气N沉降是近一、二十年来人们非常关注的全球变化现象,它们所带来的一系列生态问题已成为全球变化研究的重要议题。它们不仅影响地上植被生长和群落组成,还直接或间接地影响土壤微生物过程,而土壤微生物对此做出的响应正是生态系统反馈过程中非常重要的环节。该文分别从气候变化对土壤微生物的影响(土壤微生物量、微生物活动和微生物群落结构)和土壤微生物对气候变化的响应(凋落物分解、养分利用与循环以及养分的固持与流失)两个角度,综述近期土壤微生物对气候变暖和大气N沉降响应与适应的研究进展。气候变暖和大气N沉降对土壤微生物的影响更多地反映在微生物群落的结构和功能上,而土壤微生物量、微生物活动和群落结构的变化又会通过改变凋落物分解、养分利用和C、N循环等重要的土壤生态系统功能和过程做出响应,形成正向或负向反馈,加强或削弱气候变化给整个陆地生态系统带来的影响。然而,到目前为止土壤微生物的响应对陆地生态系统产生的最终结果仍是未决的关键性问题。     

Tree-ring growth shows that the significant population decline in Norway began decades before the Black Death

The Black Death (1349–1350 in Norway) is often cited as the cause of a severe population decline and building hiatus in the middle of the 14th century. This paper analyses this hypothesis by matching the Black Death with human and environmental impacts on tree-ring growth. The number of buildings dated by dendrochronology in Norway shows a dramatic decline several decades before the plague. In Norway, the building hiatus, which has parallels in several other places in Europe, dates from the late-13th century almost to the 16th century. The first dated houses built after the plague date from the 15th century and many of the logs have exceptionally wide tree rings compared to timber from other periods. Assuming the rapid growth was because of an open landscape, the trees are likely to have grown on infields of farms abandoned due to the 14th century population decline. Since many of these fast-growing trees germinated in the early-14th century and the number of dated buildings drops dramatically several decades before the plague, the Black Death can hardly be the only reason for the population decline in Norway and one plausible explanation is that some environmental impact occurred decades earlier. The dendroclimatological evidence of cold and wet summers in the years before the plague is suggestive, but historical sources also pinpoint famine due to crop failure. They also tell of farms being abandoned several decades before the plague and mention periods of heavy rainfall and famine in the early-14th century.     

Terrestrial nitrogen–carbon cycle interactions at the global scale

Interactions between the terrestrial nitrogen (N) and carbon (C) cycles shape the response of ecosystems to global change. However, the global distribution of nitrogen availability and its importance in global biogeochemistry and biogeochemical interactions with the climate system remain uncertain. Based on projections of a terrestrial biosphere model scaling ecological understanding of nitrogen–carbon cycle interactions to global scales, anthropogenic nitrogen additions since 1860 are estimated to have enriched the terrestrial biosphere by 1.3 Pg N, supporting the sequestration of 11.2 Pg C. Over the same time period, CO₂ fertilization has increased terrestrial carbon storage by 134.0 Pg C, increasing the terrestrial nitrogen stock by 1.2 Pg N. In 2001–2010, terrestrial ecosystems sequestered an estimated total of 27 Tg N yr⁻¹ (1.9 Pg C yr⁻¹), of which 10 Tg N yr⁻¹ (0.2 Pg C yr⁻¹) are due to anthropogenic nitrogen deposition. Nitrogen availability already limits terrestrial carbon sequestration in the boreal and temperate zone, and will constrain future carbon sequestration in response to CO₂ fertilization (regionally by up to 70% compared with an estimate without considering nitrogen–carbon interactions). This reduced terrestrial carbon uptake will probably dominate the role of the terrestrial nitrogen cycle in the climate system, as it accelerates the accumulation of anthropogenic CO₂ in the atmosphere. However, increases of N₂O emissions owing to anthropogenic nitrogen and climate change (at a rate of approx. 0.5 Tg N yr⁻¹ per 1°C degree climate warming) will add an important long-term climate forcing.     

Ensemble projections elucidate effects of uncertainty in terrestrial nitrogen limitation on future carbon uptake

The magnitude of the nitrogen (N) limitation of terrestrial carbon (C) storage over the 21st century is highly uncertain because of the complex interactions between the terrestrial C and N cycles. We use an ensemble approach to quantify and attribute process‐level uncertainty in C‐cycle projections by analysing a 30‐member ensemble representing published alternative representations of key N cycle processes (stoichiometry, biological nitrogen fixation (BNF) and ecosystem N losses) within the framework of one terrestrial biosphere model. Despite large differences in the simulated present‐day N cycle, primarily affecting simulated productivity north of 40°N, ensemble members generally conform with global C‐cycle benchmarks for present‐day conditions. Ensemble projections for two representative concentration pathways (RCP 2.6 and RCP 8.5) show that the increase in land C storage due to CO₂ fertilization is reduced by 24 ± 15% due to N constraints, whereas terrestrial C losses associated with climate change are attenuated by 19 ± 20%. As a result, N cycling reduces projected land C uptake for the years 2006–2099 by 19% (37% decrease to 3% increase) for RCP 2.6, and by 21% (40% decrease to 9% increase) for RCP 8.5. Most of the ensemble spread results from uncertainty in temperate and boreal forests, and is dominated by uncertainty in BNF (10% decrease to 50% increase for RCP 2.6, 5% decrease to 100% increase for RCP 8.5). However, choices about the flexibility of ecosystem C:N ratios and processes controlling ecosystem N losses regionally also play important roles. The findings of this study demonstrate clearly the need for an ensemble approach to quantify likely future terrestrial C–N cycle trajectories. Present‐day C‐cycle observations only weakly constrain the future ensemble spread, highlighting the need for better observational constraints on large‐scale N cycling, and N cycle process responses to global change.     

A Synthesis of Climate and Vegetation Cover Effects on Biogeochemical Cycling in Shrub-Dominated Drylands

Semi-arid and arid ecosystems dominated by shrubs (“dry shrublands”) are an important component of the global C cycle, but impacts of climate change and elevated atmospheric CO₂ on biogeochemical cycling in these ecosystems have not been synthetically assessed. This study synthesizes data from manipulative studies and from studies contrasting ecosystem processes in different vegetation microsites (that is, shrub or herbaceous canopy versus intercanopy microsites), to assess how changes in climate and atmospheric CO₂ affect biogeochemical cycles by altering plant and microbial physiology and ecosystem structure. Further, we explore how ecosystem structure impacts on biogeochemical cycles differ across a climate gradient. We found that: (1) our ability to project ecological responses to changes in climate and atmospheric CO₂ is limited by a dearth of manipulative studies, and by a lack of measurements in those studies that can explain biogeochemical changes, (2) changes in ecosystem structure will impact biogeochemical cycling, with decreasing pools and fluxes of C and N if vegetation canopy microsites were to decline, and (3) differences in biogeochemical cycling between microsites are predictable with a simple aridity index (MAP/MAT), where the relative difference in pools and fluxes of C and N between vegetation canopy and intercanopy microsites is positively correlated with aridity. We conclude that if climate change alters ecosystem structure, it will strongly impact biogeochemical cycles, with increasing aridity leading to greater heterogeneity in biogeochemical cycling among microsites. Additional long-term manipulative experiments situated across dry shrublands are required to better predict climate change impacts on biogeochemical cycling in deserts.     

Isotopic evidence for increased carbon and nitrogen exchanges between peatland plants and their symbiotic microbes with rising atmospheric CO2 concentrations since 15,000 cal. year BP

Whether nitrogen (N) availability will limit plant growth and removal of atmospheric CO₂ by the terrestrial biosphere this century is controversial. Studies have suggested that N could progressively limit plant growth, as trees and soils accumulate N in slowly cycling biomass pools in response to increases in carbon sequestration. However, a question remains over whether longer-term (decadal to century) feedbacks between climate, CO₂ and plant N uptake could emerge to reduce ecosystem-level N limitations. The symbioses between plants and microbes can help plants to acquire N from the soil or from the atmosphere via biological N₂ fixation—the pathway through which N can be rapidly brought into ecosystems and thereby partially or completely alleviate N limitation on plant productivity. Here we present measurements of plant N isotope composition (δ¹⁵N) in a peat core that dates to 15,000 cal. year BP to ascertain ecosystem-level N cycling responses to rising atmospheric CO₂ concentrations. We find that pre-industrial increases in global atmospheric CO₂ concentrations corresponded with a decrease in the δ¹⁵N of both Sphagnum moss and Ericaceae when constrained for climatic factors. A modern experiment demonstrates that the δ¹⁵N of Sphagnum decreases with increasing N₂-fixation rates. These findings suggest that plant-microbe symbioses that facilitate N acquisition are, over the long term, enhanced under rising atmospheric CO₂ concentrations, highlighting an ecosystem-level feedback mechanism whereby N constraints on terrestrial carbon storage can be overcome.     

Changes in phenological events in response to a global warming scenario reveal greater adaptability of winter annual compared with summer annual arabidopsis ecotypes

Background and AimsThe impact of global warming on life cycle timing is uncertain. We investigated changes in life cycle timing in a global warming scenario. We compared Arabidopsis thaliana ecotypes adapted to the warm/dry Cape Verdi Islands (Cvi), Macaronesia, and the cool/wet climate of the Burren (Bur), Ireland, Northern Europe. These are obligate winter and summer annuals, respectively.MethodsUsing a global warming scenario predicting a 4 °C temperature rise from 2011 to approx. 2080, we produced F₁ seeds at each end of a thermogradient tunnel. Each F₁ cohort (cool and warm) then produced F₂ seeds at both ends of the thermal gradient in winter and summer annual life cycles. F₂ seeds from the winter life cycle were buried at three positions along the gradient to determine the impact of temperature on seedling emergence in a simulated winter life cycle.Key ResultsIn a winter life cycle, increasing temperatures advanced flowering time by 10.1 d °C^–1 in the winter annual and 4.9 d °C^–1 in the summer annual. Plant size and seed yield responded positively to global warming in both ecotypes. In a winter life cycle, the impact of increasing temperature on seedling emergence timing was positive in the winter annual, but negative in the summer annual. Global warming reduced summer annual plant size and seed yield in a summer life cycle.ConclusionsSeedling emergence timing observed in the north European summer annual ecotype may exacerbate the negative impact of predicted increased spring and summer temperatures on their establishment and reproductive performance. In contrast, seedling establishment of the Macaronesian winter annual may benefit from higher soil temperatures that will delay emergence until autumn, but which also facilitates earlier spring flowering and consequent avoidance of high summer temperatures. Such plasticity gives winter annual arabidopsis ecotypes a distinct advantage over summer annuals in expected global warming scenarios. This highlights the importance of variation in the timing of seedling establishment in understanding plant species responses to anthropogenic climate change.     

Are there links between responses of soil microbes and ecosystem functioning to elevated CO2, N deposition and warming? A global perspective

In recent years, there has been an increase in research to understand how global changes’ impacts on soil biota translate into altered ecosystem functioning. However, results vary between global change effects, soil taxa, and ecosystem processes studied, and a synthesis of relationships is lacking. Therefore, here we initiate such a synthesis to assess whether the effect size of global change drivers (elevated CO₂, N deposition, and warming) on soil microbial abundance is related with the effect size of these drivers on ecosystem functioning (plant biomass, soil C cycle, and soil N cycle) using meta‐analysis and structural equation modeling. For N deposition and warming, the global change effect size on soil microbes was positively associated with the global change effect size on ecosystem functioning, and these relationships were consistent across taxa and ecosystem processes. However, for elevated CO₂, such links were more taxon and ecosystem process specific. For example, fungal abundance responses to elevated CO₂ were positively correlated with those of plant biomass but negatively with those of the N cycle. Our results go beyond previous assessments of the sensitivity of soil microbes and ecosystem processes to global change, and demonstrate the existence of general links between the responses of soil microbial abundance and ecosystem functioning. Further we identify critical areas for future research, specifically altered precipitation, soil fauna, soil community composition, and litter decomposition, that are need to better quantify the ecosystem consequences of global change impacts on soil biodiversity.     

Evolutionary History and Novel Biotic Interactions Determine Plant Responses to Elevated CO2 and Nitrogen Fertilization

A major frontier in global change research is predicting how multiple agents of global change will alter plant productivity, a critical component of the carbon cycle. Recent research has shown that plant responses to climate change are phylogenetically conserved such that species within some lineages are more productive than those within other lineages in changing environments. However, it remains unclear how phylogenetic patterns in plant responses to changing abiotic conditions may be altered by another agent of global change, the introduction of non-native species. Using a system of 28 native Tasmanian Eucalyptus species belonging to two subgenera, Symphyomyrtus and Eucalyptus, we hypothesized that productivity responses to abiotic agents of global change (elevated CO₂ and increased soil N) are unique to lineages, but that novel interactions with a non-native species mediate these responses. We tested this hypothesis by examining productivity of 1) native species monocultures and 2) mixtures of native species with an introduced hardwood plantation species, Eucalyptus nitens, to experimentally manipulated soil N and atmospheric CO₂. Consistent with past research, we found that N limits productivity overall, especially in elevated CO₂ conditions. However, monocultures of species within the Symphyomyrtus subgenus showed the strongest response to N (gained 127% more total biomass) in elevated CO₂ conditions, whereas those within the Eucalyptus subgenus did not respond to N. Root:shoot ratio (an indicator of resource use) was on average greater in species pairs containing Symphyomyrtus species, suggesting that functional traits important for resource uptake are phylogenetically conserved and explaining the phylogenetic pattern in plant response to changing environmental conditions. Yet, native species mixtures with E. nitens exhibited responses to CO₂ and N that differed from those of monocultures, supporting our hypothesis and highlighting that both plant evolutionary history and introduced species will shape community productivity in a changing world.     

Do dams and levees impact nitrogen cycling? Simulating the effects of flood alterations on floodplain denitrification

A fundamental challenge in understanding the global nitrogen cycle is the quantification of denitrification on large heterogeneous landscapes. Because floodplains are important sites for denitrification and nitrogen retention, we developed a generalized floodplain biogeochemical model to determine whether dams and flood‐control levees affect floodplain denitrification by altering floodplain inundation. We combined a statistical model of floodplain topography with a model of hydrology and nitrogen biogeochemistry to simulate floods of different magnitude. The model predicted substantial decreases in NO₃‐N processing on floodplains whose overbank floods have been altered by levees and upstream dams. Our simulations suggest that dams may reduce nitrate processing more than setback levees. Levees increased areal floodplain denitrification rates, but this effect was offset by a reduction in the area inundated. Scenarios that involved a levee also resulted in more variability in N processing among replicate floodplains. Nitrate loss occurred rapidly and completely in our model floodplains. As a consequence, total flood volume and the initial mass of nitrate reaching a floodplain may provide reasonable estimates of total N processing on floodplains during floods. This finding suggests that quantifying the impact of dams and levees on floodplain denitrification may be possible using recent advances in remote sensing of floodplain topography and flood stage. Furthermore, when considering flooding over the long‐term, the cumulative N processed by frequent smaller floods was estimated to be quite large relative to that processed by larger, less frequent floods. Our results suggest that floodplain denitrification may be greatly influenced by the pervasive anthropogenic flood‐control measures that currently exist on most majors river floodplains throughout the world, and may have the potential to be impacted by future changes in flood probabilities that will likely occur as a result of climate shifts.     

Nitrogen deposition and greenhouse gas emissions from grasslands: uncertainties and future directions

Increases in atmospheric nitrogen deposition (N_dep) can strongly affect the greenhouse gas (GHG; CO₂, CH₄, and N₂O) sink capacity of grasslands as well as other terrestrial ecosystems. Robust predictions of the net GHG sink strength of grasslands depend on how experimental N loads compare to projected N_dep rates, and how accurately the relationship between GHG fluxes and N_dep is characterized. A literature review revealed that the vast majority of experimental N loads were higher than levels these ecosystems are predicted to experience in the future. Using a process‐based biogeochemical model, we predicted that low levels of N_dep either enhanced or reduced the net GHG sink strength of most grasslands, but as experimental N loads continued to increase, grasslands transitioned to a N saturation‐decline stage, where the sensitivity of GHG exchange to further increases in N_dep declined. Most published studies represented treatments well into the N saturation‐decline stage. Our model results predict that the responses of GHG fluxes to N are highly nonlinear and that the N saturation thresholds for GHGs varied greatly among grasslands and with fire management. We predict that during the 21st century some grasslands will be in the N limitation stage where others will transition into the N saturation‐decline stage. The linear relationship between GHG sink strength and N load assumed by most studies can overestimate or underestimate predictions of the net GHG sink strength of grasslands depending on their N baseline status. The next generation of global change experiments should be designed at multiple N loads consistent with future N_dep rates to improve our empirical understanding and predictive ability.     

Influences of infrequent fire, elevation and pre-fire vegetation on the persistence of quaking aspen (Populus tremuloides Michx.) in the Flat Tops area, Colorado, USA

Aim  The recent concern that quaking aspen ( Populus tremuloides Michx.) has been declining in parts of western North America due to fire suppression is largely based on trends during the latter part of the 20th century. The aim of the current study was to compare the extent of aspen in the modern landscape with its extent in the late 19th century prior to fire suppression, and to assess the effects of elevation, late-19th century fires, and pre-fire forest composition on the successional status of aspen.
Location  North-west Colorado, USA.
Methods  We used a georeferenced 1898 map and modern maps to examine trends in aspen dominance since the late 19th century in a 348,586 ha area of White River and Routt National Forests in north-western Colorado. Stand age and structure were sampled in 30 stands.
Results  We found no evidence of overall aspen decline over this period. In fact, aspen distribution has increased in parts of the study area following severe fires in the late 19th century in forests that were previously dominated by conifers. Aspen persistence and increase was especially pronounced at elevations below 3000 m a.s.l. Most 120-year-old post-fire stands that are presently being successionally replaced by conifers were dominated by conifers prior to the last severe fire.
Main conclusions  Human perceptions of ecosystems are often on time scales that are shorter than the cycles of natural variation within those ecosystems. This disparity may lead to an underestimation of the range of natural variability of ecosystem patterns and processes. The appropriate temporal scale of inquiry is necessary for the correct understanding of natural variation in ecosystems.