Decadal-scale Dynamics of Water, Carbon and Nitrogen in a California Chaparral Ecosystem: DAYCENT Modeling Results

The Mediterranean climate, with its characteristic of dry summers and wet winters, influences the hydrologic and microbial processes that control carbon (C) and nitrogen (N) biogeochemical processes in chaparral ecosystems. These biogeochemical processes in turn determine N cycling under chronic N deposition. In order to examine connections between climate and N dynamics, we quantified decadal-scale water, C and N states and fluxes at annual, monthly and daily time steps for a California chaparral ecosystem in the Sierra Nevada using the DAYCENT model. The daily output simulations of net mineralization, stream flow and stream nitrate (NO₃⁻) export were developed for DAYCENT in order to simulate the N dynamics most appropriate for the abrupt rewetting events characteristic of Mediterranean chaparral ecosystems. Overall, the magnitude of annual modeled net N mineralization, soil and plant biomass C and N, nitrate export and gaseous N emission agreed with those of observations. Gaseous N emission was a major N loss pathway in chaparral ecosystems, in which nitric oxide (NO) is the dominant species. The modeled C and N fluxes of net primary production (NPP), N uptake and N mineralization, NO₃⁻ export and gaseous N emission showed both high inter-annual and intra-annual variability. Our simulations also showed dramatic fire effects on NPP, N uptake, N mineralization and gaseous N emission for three years of postfire. The decease in simulated soil organic C and N storages was not dramatic, but lasted a longer time. For the seasonal pattern, the predicted C and N fluxes were greatest during December to March, and lowest in the summer. The model predictions suggested that an increase in the N deposition rate would increase N losses through gaseous N emission and stream N export in the chaparral ecosystems of the Sierra Nevada due to changes in N saturation status. The model predictions could not capture stream NO₃⁻ export during most rewetting events suggesting that a dry-rewetting mechanism representing the increase in N mineralization following soil wetting needs to be incorporated into biogeochemical models of semi-arid ecosystems.     

Carbon cost of plant nitrogen acquisition: global carbon cycle impact from an improved plant nitrogen cycle in the Community Land Model

Plants typically expend a significant portion of their available carbon (C) on nutrient acquisition – C that could otherwise support growth. However, given that most global terrestrial biosphere models (TBMs) do not include the C cost of nutrient acquisition, these models fail to represent current and future constraints to the land C sink. Here, we integrated a plant productivity‐optimized nutrient acquisition model – the Fixation and Uptake of Nitrogen Model – into one of the most widely used TBMs, the Community Land Model. Global plant nitrogen (N) uptake is dynamically simulated in the coupled model based on the C costs of N acquisition from mycorrhizal roots, nonmycorrhizal roots, N‐fixing microbes, and retranslocation (from senescing leaves). We find that at the global scale, plants spend 2.4 Pg C yr^?1 to acquire 1.0 Pg N yr^?1, and that the C cost of N acquisition leads to a downregulation of global net primary production (NPP) by 13%. Mycorrhizal uptake represented the dominant pathway by which N is acquired, accounting for ~66% of the N uptake by plants. Notably, roots associating with arbuscular mycorrhizal (AM) fungi – generally considered for their role in phosphorus (P) acquisition – are estimated to be the primary source of global plant N uptake owing to the dominance of AM‐associated plants in mid‐ and low‐latitude biomes. Overall, our coupled model improves the representations of NPP downregulation globally and generates spatially explicit patterns of belowground C allocation, soil N uptake, and N retranslocation at the global scale. Such model improvements are critical for predicting how plant responses to altered N availability (owing to N deposition, rising atmospheric CO₂, and warming temperatures) may impact the land C sink.     

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.     

Global‐scale impacts of nitrogen deposition on tree carbon sequestration in tropical,temperate, and boreal forests: A meta‐analysis

Elevated nitrogen (N) deposition may increase net primary productivity in N‐limited terrestrial ecosystems and thus enhance the terrestrial carbon (C) sink. To assess the magnitude of this N‐induced C sink, we performed a meta‐analysis on data from forest fertilization experiments to estimate N‐induced C sequestration in aboveground tree woody biomass, a stable C pool with long turnover times. Our results show that boreal and temperate forests responded strongly to N addition and sequestered on average an additional 14 and 13 kg C per kg N in aboveground woody biomass, respectively. Tropical forests, however, did not respond significantly to N addition. The common hypothesis that tropical forests do not respond to N because they are phosphorus‐limited could not be confirmed, as we found no significant response to phosphorus addition in tropical forests. Across climate zones, we found that young forests responded more strongly to N addition, which is important as many previous meta‐analyses of N addition experiments rely heavily on data from experiments on seedlings and young trees. Furthermore, the C–N response (defined as additional mass unit of C sequestered per additional mass unit of N addition) was affected by forest productivity, experimental N addition rate, and rate of ambient N deposition. The estimated C–N responses from our meta‐analysis were generally lower that those derived with stoichiometric scaling, dynamic global vegetation models, and forest growth inventories along N deposition gradients. We estimated N‐induced global C sequestration in tree aboveground woody biomass by multiplying the C–N responses obtained from the meta‐analysis with N deposition estimates per biome. We thus derived an N‐induced global C sink of about 177 (112–243) Tg C/year in aboveground and belowground woody biomass, which would account for about 12% of the forest biomass C sink (1,400 Tg C/year).     

The impact of elevated CO2, increased nitrogen availability and biodiversity on plant tissue quality and decomposition

Elevated CO₂, increased nitrogen (N) deposition and increasing species richness can increase net primary productivity (NPP). However, unless there are comparable changes in decomposition, increases in productivity will most likely be unsustainable. Without comparable increases in decomposition nutrients would accumulate in dead organic matter leading to nutrient limitations that could eventually prohibit additional increases in productivity. To address this issue, we measured aboveground plant and litter quality and belowground root quality, as well as decomposition of aboveground litter for one and 2‐year periods using in situ litterbags in response to a three‐way factorial manipulation of CO₂ (ambient vs. 560 ppm), N deposition (ambient vs. the addition of 4 g N m⁻² yr⁻¹) and plant species richness (one, four, nine and 16 species) in experimental grassland plots. Litter chemistry responded to the CO₂, N and plant diversity treatments, but decomposition was much less responsive. Elevated CO₂ induced decreases in % N and % lignin in plant tissues. N addition led to increases in % N and decreases in % lignin. Increasing plant diversity led to decreases in % N and % lignin and an increase in % cellulose. In contrast to the litter chemistry changes, elevated CO₂ had a much lower impact on decomposition and resulted in only a 2.5% decrease in carbon (C) loss. Detectable responses were not observed either to N addition or to species richness. These results suggest that global change factors such as biodiversity loss, elevated CO₂ and N deposition lead to significant changes in tissue quality; however, the response of decomposition is modest. Thus, the observed increases in productivity at higher diversity levels and with elevated CO₂ and N fertilization are not matched by an increase in decomposition rates. This lack of coupled responses between production and decomposition is likely to result in an accumulation of nutrients in the litter pool which will dampen the response of NPP to these factors over time.     

Net Primary Production and Carbon Stocks for Subarctic Mesic–Dry Tundras with Contrasting Microtopography, Altitude, and Dominant Species

Mesic–dry tundras are widespread in the Arctic but detailed assessments of net primary production (NPP) and ecosystem carbon (C) stocks are lacking. We addressed this lack of knowledge by determining the seasonal dynamics of aboveground vascular NPP, annual NPP, and whole-ecosystem C stocks in five mesic–dry tundras in Northern Sweden with contrasting microtopography, altitude, and dominant species. Those measurements were paralleled by the stock assessments of nitrogen (N), the limiting nutrient. The vascular production was determined by harvest or in situ growing units, whereas the nonvascular production was obtained from average species growth rates, previously assessed at the sites. Results showed that aboveground vascular NPP (15–270 g m⁻²), annual NPP (214–282 g m⁻² or 102–137 g C m⁻²) and vegetation biomass (330–2450 g m⁻²) varied greatly among communities. Vegetation dominated by Empetrum hermaphroditum is more productive than Cassiope tetragona vegetation. Although the large majority of the apical NPP occurred in early-mid season (85%), production of stems and evergreen leaves proceeded until about 2 weeks before senescence. Most of the vascular vegetation was belowground (80%), whereas most of the vegetation production occurred aboveground (85%). Ecosystem C and N stocks were 2100–8200 g C m⁻² and 80–330 g N m⁻², respectively, stored mainly in the soil turf and in the fine organic soil. Such stocks are comparable to the C and N stocks of moister tundra types, such as tussock tundra. Author Contributions  Matteo Campioli, Anders Michelsen, Roeland Samson, Raoul Lemeur—conceived and designed study, Matteo Campioli, Anders Michelsen, Andreas Demey, Annemie Vermeulen—performed research, Matteo Campioli—analyzed data, and Matteo Campioli—wrote the paper.     

Long‐term nitrogen addition modifies microbial composition and functions for slow carbon cycling and increased sequestration in tropical forest soil

Nitrogen (N) deposition is a component of global change that has considerable impact on belowground carbon (C) dynamics. Plant growth stimulation and alterations of fungal community composition and functions are the main mechanisms driving soil C gains following N deposition in N‐limited temperate forests. In N‐rich tropical forests, however, N deposition generally has minor effects on plant growth; consequently, C storage in soil may strongly depend on the microbial processes that drive litter and soil organic matter decomposition. Here, we investigated how microbial functions in old‐growth tropical forest soil responded to 13 years of N addition at four rates: 0 (Control), 50 (Low‐N), 100 (Medium‐N), and 150 (High‐N) kg N ha^?1 year^?1. Soil organic carbon (SOC) content increased under High‐N, corresponding to a 33% decrease in CO₂ efflux, and reductions in relative abundances of bacteria as well as genes responsible for cellulose and chitin degradation. A 113% increase in N₂O emission was positively correlated with soil acidification and an increase in the relative abundances of denitrification genes (narG and norB). Soil acidification induced by N addition decreased available P concentrations, and was associated with reductions in the relative abundance of phytase. The decreased relative abundance of bacteria and key functional gene groups for C degradation were related to slower SOC decomposition, indicating the key mechanisms driving SOC accumulation in the tropical forest soil subjected to High‐N addition. However, changes in microbial functional groups associated with N and P cycling led to coincidentally large increases in N₂O emissions, and exacerbated soil P deficiency. These two factors partially offset the perceived beneficial effects of N addition on SOC storage in tropical forest soils. These findings suggest a potential to incorporate microbial community and functions into Earth system models considering their effects on greenhouse gas emission, biogeochemical processes, and biodiversity of tropical ecosystems.     

Reduced N cycling in response to elevated CO2, warming,and drought in a Danish heathland: Synthesizing results of the CLIMAITE project after two years of treatments

Field‐scale experiments simulating realistic future climate scenarios are important tools for investigating the effects of current and future climate changes on ecosystem functioning and biogeochemical cycling. We exposed a seminatural Danish heathland ecosystem to elevated atmospheric carbon dioxide (CO₂), warming, and extended summer drought in all combinations. Here, we report on the short‐term responses of the nitrogen (N) cycle after 2 years of treatments. Elevated CO₂ significantly affected aboveground stoichiometry by increasing the carbon to nitrogen (C/N) ratios in the leaves of both co‐dominant species (Calluna vulgaris and Deschampsia flexuosa), as well as the C/N ratios of Calluna flowers and by reducing the N concentration of Deschampsia litter. Belowground, elevated CO₂ had only minor effects, whereas warming increased N turnover, as indicated by increased rates of microbial NH₄⁺ consumption, gross mineralization, potential nitrification, denitrification and N₂O emissions. Drought reduced belowground gross N mineralization and decreased fauna N mass and fauna N mineralization. Leaching was unaffected by treatments but was significantly higher across all treatments in the second year than in the much drier first year indicating that ecosystem N loss is highly sensitive to changes and variability in amount and timing of precipitation. Interactions between treatments were common and although some synergistic effects were observed, antagonism dominated the interactive responses in treatment combinations, i.e. responses were smaller in combinations than in single treatments. Nonetheless, increased C/N ratios of photosynthetic tissue in response to elevated CO₂, as well as drought‐induced decreases in litter N production and fauna N mineralization prevailed in the full treatment combination. Overall, the simulated future climate scenario therefore lead to reduced N turnover, which could act to reduce the potential growth response of plants to elevated atmospheric CO₂ concentration.     

A historical meta‐analysis of global terrestrial net primary productivity: are estimates converging?

Net primary productivity (NPP) is one of the most important ecosystem parameters, representing vegetation activity, biogeochemical cycling, and ecosystem services. To assess how well the scientific community understands the biospheric function, a historical meta‐analysis was conducted. By surveying the literature from 1862 to 2011, I extracted 251 estimates of total terrestrial NPP at the present time (NPP_T) and calculated their statistical metrics. For all the data, the mean±standard deviation and median were 56.2±14.3 and 56.4 Pg C yr^–1, respectively. Even for estimates published after 2000, a substantial level of uncertainty (coefficient of variation by ±15%) was inevitable. The estimates were categorized on the basis of methodology (i.e., inventory analysis, empirical model, biogeochemical model, dynamic global vegetation model, and remote sensing) to examine the consistency among the statistical metrics of each category. Chronological analysis revealed that the present NPP_T estimates were directed by extensive field surveys in the 1960s and 1970s (e.g., the International Biological Programme). A wide range of uncertainty remains in modern estimates based on advanced biogeochemical and dynamic vegetation models and remote‐sensing techniques. Several critical factors accounting for the estimation uncertainty are discussed. Ancillary analyses were performed to derive additional ecological and human‐related parameters related to NPP. For example, interannual variability, carbon‐use efficiency (a ratio of NPP to gross photosynthesis), human appropriation, and preindustrial NPP_T were assessed. Finally, I discuss the importance of improving NPP_T estimates in the context of current global change studies and integrated carbon cycle research.     

Net greenhouse gas balance in response to nitrogen enrichment: perspectives from a coupled biogeochemical model

Increasing reactive nitrogen (N) input has been recognized as one of the important factors influencing climate system through affecting the uptake and emission of greenhouse gases (GHG). However, the magnitude and spatiotemporal variations of N‐induced GHG fluxes at regional and global scales remain far from certain. Here we selected China as an example, and used a coupled biogeochemical model in conjunction with spatially explicit data sets (including climate, atmospheric CO₂, O₃, N deposition, land use, and land cover changes, and N fertilizer application) to simulate the concurrent impacts of increasing atmospheric and fertilized N inputs on balance of three major GHGs (CO₂, CH₄, and N₂O). Our simulations showed that these two N enrichment sources in China decreased global warming potential (GWP) through stimulating CO₂ sink and suppressing CH₄ emission. However, direct N₂O emission was estimated to offset 39% of N‐induced carbon (C) benefit, with a net GWP of three GHGs averaging ?376.3 ± 146.4 Tg CO₂ eq yr^?1 (the standard deviation is interannual variability of GWP) during 2000–2008. The chemical N fertilizer uses were estimated to increase GWP by 45.6 ± 34.3 Tg CO₂ eq yr^?1 in the same period, and C sink was offset by 136%. The largest C sink offset ratio due to increasing N input was found in Southeast and Central mainland of China, where rapid industrial development and intensively managed crop system are located. Although exposed to the rapidly increasing N deposition, most of the natural vegetation covers were still showing decreasing GWP. However, due to extensive overuse of N fertilizer, China's cropland was found to show the least negative GWP, or even positive GWP in recent decade. From both scientific and policy perspectives, it is essential to incorporate multiple GHGs into a coupled biogeochemical framework for fully assessing N impacts on climate changes.     

Experimental fire increases soil carbon dioxide efflux in a grassland long‐term multifactor global change experiment

Numerous studies have demonstrated that soil respiration rates increase under experimental warming, although the long‐term, multiyear dynamics of this feedback are not well constrained. Less is known about the effects of single, punctuated events in combination with other longer‐duration anthropogenic influences on the dynamics of soil carbon (C) loss. In 2012 and 2013, we assessed the effects of decadal‐scale anthropogenic global change – warming, increased nitrogen (N) deposition, elevated carbon dioxide (CO₂), and increased precipitation – on soil respiration rates in an annual‐dominated Mediterranean grassland. We also investigated how controlled fire and an artificial wet‐up event, in combination with exposure to the longer‐duration anthropogenic global change factors, influenced the dynamics of C cycling in this system. Decade‐duration surface soil warming (1–2 °C) had no effect on soil respiration rates, while +N addition and elevated CO₂ concentrations increased growing‐season soil CO₂ efflux rates by increasing annual aboveground net primary production (NPP) and belowground fine root production, respectively. Low‐intensity experimental fire significantly elevated soil CO₂ efflux rates in the next growing season. Based on mixed‐effects modeling and structural equation modeling, low‐intensity fire increased growing‐season soil respiration rates through a combination of three mechanisms: large increases in soil temperature (3–5 °C), significant increases in fine root production, and elevated aboveground NPP. Our study shows that in ecosystems where soil respiration has acclimated to moderate warming, further increases in soil temperature can stimulate greater soil CO₂ efflux. We also demonstrate that punctuated short‐duration events such as fire can influence soil C dynamics with implications for both the parameterization of earth system models (ESMs) and the implementation of climate change mitigation policies that involve land‐sector C accounting.     

Elevated carbon dioxide and ozone alter productivity and ecosystem carbon content in northern temperate forests

Three young northern temperate forest communities in the north‐central United States were exposed to factorial combinations of elevated carbon dioxide (CO₂) and tropospheric ozone (O₃) for 11 years. Here, we report results from an extensive sampling of plant biomass and soil conducted at the conclusion of the experiment that enabled us to estimate ecosystem carbon (C) content and cumulative net primary productivity (NPP). Elevated CO₂ enhanced ecosystem C content by 11%, whereas elevated O₃ decreased ecosystem C content by 9%. There was little variation in treatment effects on C content across communities and no meaningful interactions between CO₂ and O₃. Treatment effects on ecosystem C content resulted primarily from changes in the near‐surface mineral soil and tree C, particularly differences in woody tissues. Excluding the mineral soil, cumulative NPP was a strong predictor of ecosystem C content (r² = 0.96). Elevated CO₂ enhanced cumulative NPP by 39%, a consequence of a 28% increase in canopy nitrogen (N) content (g N m^?2) and a 28% increase in N productivity (NPP/canopy N). In contrast, elevated O₃ lowered NPP by 10% because of a 21% decrease in canopy N, but did not impact N productivity. Consequently, as the marginal impact of canopy N on NPP (?NPP/?N) decreased through time with further canopy development, the O₃ effect on NPP dissipated. Within the mineral soil, there was less C in the top 0.1 m of soil under elevated O₃ and less soil C from 0.1 to 0.2 m in depth under elevated CO₂. Overall, these results suggest that elevated CO₂ may create a sustained increase in NPP, whereas the long‐term effect of elevated O₃ on NPP will be smaller than expected. However, changes in soil C are not well‐understood and limit our ability to predict changes in ecosystem C content.     

Divergence in plant and microbial allocation strategies explains continental patterns in microbial allocation and biogeochemical fluxes

Allocation trade‐offs shape ecological and biogeochemical phenomena at local to global scale. Plant allocation strategies drive major changes in ecosystem carbon cycling. Microbial allocation to enzymes that decompose carbon vs. organic nutrients may similarly affect ecosystem carbon cycling. Current solutions to this allocation problem prioritise stoichiometric tradeoffs implemented in plant ecology. These solutions may not maximise microbial growth and fitness under all conditions, because organic nutrients are also a significant carbon resource for microbes. I created multiple allocation frameworks and simulated microbial growth using a microbial explicit biogeochemical model. I demonstrate that prioritising stoichiometric trade‐offs does not optimise microbial allocation, while exploiting organic nutrients as carbon resources does. Analysis of continental‐scale enzyme data supports the allocation patterns predicted by this framework, and modelling suggests large deviations in soil C loss based on which strategy is implemented. Therefore, understanding microbial allocation strategies will likely improve our understanding of carbon cycling and climate.     

Primary production and carbon allocation in relation to nutrient supply in a tropical experimental forest

Nutrient supply commonly limits aboveground plant productivity in forests, but the effects of an altered nutrient supply on gross primary production (GPP) and patterns of carbon (C) allocation remain poorly characterized. Increased nutrient supply may lead to a higher aboveground net primary production (ANPP), but a lower total belowground carbon allocation (TBCA), with little change in either aboveground plant respiration (APR) or GPP. Alternatively, increases in nutrient supply may increase GPP, with the quantity of GPP allocated aboveground increasing more steeply than the quantity of GPP allocated belowground. To examine the effects of an elevated nutrient supply on the C allocation patterns in forests, we determined whole‐ecosystem C budgets in unfertilized plots of Eucalyptus saligna and in adjacent plots receiving regular additions of 65 kg N ha^?1, 31 kg P ha^?1, 46 kg K ha^?1, and macro‐ and micronutrients. We measured the absolute flux of C allocated to the components of GPP (ANPP, TBCA and APR), as well as the fraction of GPP allocated to these components. Fertilization dramatically increased GPP. Averaged over 3 years, GPP in the fertilized plots was 34% higher than that in the unfertilized controls (3.95 vs. 2.95 kg C m^?2 yr^?1). Fertilization‐related increases in GPP were allocated entirely aboveground – ANPP was 85% higher and APR was 57% higher in the fertilized than in the control plots, while TBCA did not differ significantly between treatments. Carbon use efficiency (NPP/GPP) was slightly higher in the fertilized (0.53) compared with the control plots (0.51). Overall, fertilization increased ANPP and APR, and these increases were related to a greater GPP and an increase in the fraction of GPP allocated aboveground.     

Microbial biomass and nitrogen cycling responses to fertilization and litter removal in young northern hardwood forests

The influence of site fertility on soil microbial biomass and activity is not well understood but is likely to be complex because of interactions with plant responses to nutrient availability. We examined the effects of long-term (8 yr) fertilization and litter removal on forest floor microbial biomass and N and C transformations to test the hypothesis that higher soil resource availability stimulates microbial activity. Microbial biomass and respiration decreased by 20–30 % in response to fertilization. Microbial C averaged 3.8 mg C/g soil in fertilized, 5.8 mg C/g in control, and 5.5 mg C/g in litter removal plots. Microbial respiration was 200 µg CO₂-C g^–1 d^–1 in fertilized plots, compared to 270 µg CO₂-C g^–1 d^–1 in controls. Gross N mineralization and N immobilization did not differ among treatments, despite higher litter nutrient concentrations in fertilized plots and the removal of substantial quantities of C and N in litter removal plots. Net N mineralization was significantly reduced by fertilization. Gross nitrification and NO₃ ^– immobilization both were increased by fertilization. Nitrate thus became a more important part of microbial N cycling in fertilized plots even though NH₄ ⁺ availability was not stimulated by fertilization.Soil microorganisms did not mineralize more C or N in response to fertilization and higher litter quality; instead, results suggest a difference in the physiological status of microbial biomass in fertilized plots that influenced N transformations. Respiration quotients (qCO₂, respiration per unit biomass) were higher in fertilized plots (56 µg CO₂-C mg C^–1 d^–1) than control (48 µg CO₂-C mg C^–1 d ^–1) or litter removal (45 µg CO₂-C mg C^–1 d^–1), corresponding to higher microbial growth efficiency, higher proportions of gross mineralization immobilized, and lower net N mineralization in fertilized plots. While microbial biomass is an important labile nutrient pool, patterns of microbial growth and turnover were distinct from this pool and were more important to microbial function in nitrogen cycling.     

Forest productivity under elevated CO₂ and O₃: positive feedbacks to soil N cycling sustain decade-long net primary productivity enhancement by CO₂

The accumulation of anthropogenic CO? in the Earth's atmosphere, and hence the rate of climate warming, is sensitive to stimulation of plant growth by higher concentrations of atmospheric CO?. Here, we synthesise data from a field experiment in which three developing northern forest communities have been exposed to factorial combinations of elevated CO? and O?. Enhanced net primary productivity (NPP) (c. 26% increase) under elevated CO? was sustained by greater root exploration of soil for growth-limiting N, as well as more rapid rates of litter decomposition and microbial N release during decay. Despite initial declines in forest productivity under elevated O?, compensatory growth of O? -tolerant individuals resulted in equivalent NPP under ambient and elevated O?. After a decade, NPP has remained enhanced under elevated CO? and has recovered under elevated O? by mechanisms that remain un-calibrated or not considered in coupled climate-biogeochemical models simulating interactions between the global C cycle and climate warming.     

Experimentally increased nutrient availability at the permafrost thaw front selectively enhances biomass production of deep‐rooting subarctic peatland species

Climate warming increases nitrogen (N) mineralization in superficial soil layers (the dominant rooting zone) of subarctic peatlands. Thawing and subsequent mineralization of permafrost increases plant‐available N around the thaw‐front. Because plant production in these peatlands is N‐limited, such changes may substantially affect net primary production and species composition. We aimed to identify the potential impact of increased N‐availability due to permafrost thawing on subarctic peatland plant production and species performance, relative to the impact of increased N‐availability in superficial organic layers. Therefore, we investigated whether plant roots are present at the thaw‐front (45 cm depth) and whether N‐uptake (¹⁵N‐tracer) at the thaw‐front occurs during maximum thaw‐depth, coinciding with the end of the growing season. Moreover, we performed a unique 3‐year belowground fertilization experiment with fully factorial combinations of deep‐ (thaw‐front) and shallow‐fertilization (10 cm depth) and controls. We found that certain species are present with roots at the thaw‐front (Rubus chamaemorus) and have the capacity (R. chamaemorus, Eriophorum vaginatum) for N‐uptake from the thaw‐front between autumn and spring when aboveground tissue is largely senescent. In response to 3‐year shallow‐belowground fertilization (S) both shallow‐ (Empetrum hermaphroditum) and deep‐rooting species increased aboveground biomass and N‐content, but only deep‐rooting species responded positively to enhanced nutrient supply at the thaw‐front (D). Moreover, the effects of shallow‐fertilization and thaw‐front fertilization on aboveground biomass production of the deep‐rooting species were similar in magnitude (S: 71%; D: 111% increase compared to control) and additive (S + D: 181% increase). Our results show that plant‐available N released from thawing permafrost can form a thus far overlooked additional N‐source for deep‐rooting subarctic plant species and increase their biomass production beyond the already established impact of warming‐driven enhanced shallow N‐mineralization. This may result in shifts in plant community composition and may partially counteract the increased carbon losses from thawing permafrost.     

Global variations and controlling factors of anammox rates

Soil anammox is an environmentally friendly way to eliminate reactive nitrogen (N) without generating nitrous oxide. Nevertheless, the current earth system models have not incorporated the anammox due to the lack of parameters in anammox rates on a global scale, limiting the accurate projection for N cycling. A global synthesis with 1212 observations from 89 peer-reviewed papers showed that the average anammox rate was 1.60 ± 0.17 nmol N g⁻¹ h⁻¹ in terrestrial ecosystems, with significant variations across different ecosystems. Wetlands exhibited the highest rate (2.17 ± 0.31 nmol N g⁻¹ h⁻¹), followed by croplands at 1.02 ± 0.09 nmol N g⁻¹ h⁻¹. The lowest anammox rates were observed in forests and grasslands. The anammox rates were positively correlated with the mean annual temperature, mean annual precipitation, soil moisture, organic carbon (C), total N, as well as nitrite and ammonium concentrations, but negatively with the soil C:N ratio. Structural equation models revealed that the geographical variations in anammox rates were primarily influenced by the N contents (such as nitrite and ammonium) and abundance of anammox bacteria, which collectively accounted for 42% of the observed variance. Furthermore, the abundance of anammox bacteria was well simulated by the mean annual precipitation, soil moisture, and ammonium concentrations, and 51% variance of the anammox bacteria was accounted for. The key controlling factors for soil anammox rates differed from ecosystem type, for example, organic C, total N, and ammonium contents in croplands, versus soil C:N ratio and nitrite concentrations in wetlands. The controlling factors in soil anammox rate identified by this study are useful to construct an accurate anammox module for N cycling in earth system models.     

FLUXNET and modelling the global carbon cycle

Measurements of the net CO₂ flux between terrestrial ecosystems and the atmosphere using the eddy covariance technique have the potential to underpin our interpretation of regional CO₂ source–sink patterns, CO₂ flux responses to forcings, and predictions of the future terrestrial C balance. Information contained in FLUXNET eddy covariance data has multiple uses for the development and application of global carbon models, including evaluation/validation, calibration, process parameterization, and data assimilation. This paper reviews examples of these uses, compares global estimates of the dynamics of the global carbon cycle, and suggests ways of improving the utility of such data for global carbon modelling. Net ecosystem exchange of CO₂ (NEE) predicted by different terrestrial biosphere models compares favourably with FLUXNET observations at diurnal and seasonal timescales. However, complete model validation, particularly over the full annual cycle, requires information on the balance between assimilation and decomposition processes, information not readily available for most FLUXNET sites. Site history, when known, can greatly help constrain the model‐data comparison. Flux measurements made over four vegetation types were used to calibrate the land‐surface scheme of the Goddard Institute for Space Studies global climate model, significantly improving simulated climate and demonstrating the utility of diurnal FLUXNET data for climate modelling. Land‐surface temperatures in many regions cool due to higher canopy conductances and latent heat fluxes, and the spatial distribution of CO₂ uptake provides a significant additional constraint on the realism of simulated surface fluxes. FLUXNET data are used to calibrate a global production efficiency model (PEM). This model is forced by satellite‐measured absorbed radiation and suggests that global net primary production (NPP) increased 6.2% over 1982–1999. Good agreement is found between global trends in NPP estimated by the PEM and a dynamic global vegetation model (DGVM), and between the DGVM and estimates of global NEE derived from a global inversion of atmospheric CO₂ measurements. Combining the PEM, DGVM, and inversion results suggests that CO₂ fertilization is playing a major role in current increases in NPP, with lesser impacts from increasing N deposition and growing season length. Both the PEM and the inversion identify the Amazon basin as a key region for the current net terrestrial CO₂ uptake (i.e. 33% of global NEE), as well as its interannual variability. The inversion's global NEE estimate of −1.2 Pg [C] yr⁻¹ for 1982–1995 is compatible with the PEM‐ and DGVM‐predicted trends in NPP. There is, thus, a convergence in understanding derived from process‐based models, remote‐sensing‐based observations, and inversion of atmospheric data. Future advances in field measurement techniques, including eddy covariance (particularly concerning the problem of night‐time fluxes in dense canopies and of advection or flow distortion over complex terrain), will result in improved constraints on land‐atmosphere CO₂ fluxes and the rigorous attribution of mechanisms to the current terrestrial net CO₂ uptake and its spatial and temporal heterogeneity. Global ecosystem models play a fundamental role in linking information derived from FLUXNET measurements to atmospheric CO₂ variability. A number of recommendations concerning FLUXNET data are made, including a request for more comprehensive site data (particularly historical information), more measurements in undisturbed ecosystems, and the systematic provision of error estimates. The greatest value of current FLUXNET data for global carbon cycle modelling is in evaluating process representations, rather than in providing an unbiased estimate of net CO₂ exchange.     

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.