A. Tietema

Nitrogen deposition makes a minor contribution to carbon sequestration in temperate forests

Humans have altered global nitrogen cycling such that more atmospheric N2 is being converted (‘fixed’) into biologically reactive forms by anthropogenic activities than by all natural processes combined. In particular, nitrogen oxides emitted during fuel combustion and ammonia volatilized as a result of intensive agriculture have increased atmospheric nitrogen inputs (mostly NO3 and NH4) to temperate forests in the Northern Hemisphere. Because tree growth in northern temperate regions is typically nitrogen-limited, increased nitrogen deposition could have the effect of attenuating rising atmospheric CO2 by stimulating the accumulation of forest biomass. Forest inventories indicate that the carbon contents of northern forests have increased concurrently with nitrogen deposition since the 1950s. In addition, variations in atmospheric CO2 indicate a globally significant carbon sink in northern mid-latitude forest regions. It is unclear, however, whether elevated nitrogen deposition or other factors are the primary cause of carbon sequestration in northern forests. Here we use evidence from 15N-tracer studies in nine forests to show that elevated nitrogen deposition is unlikely to be a major contributor to the putative CO2 sink in forested northern temperature regions.

Impact of nitrogen deposition on nitrogen cycling in forests: a synthesis of NITREX data

Abstract Impact of nitrogen (N) deposition was studied by comparing N fluxes, N concentrations and N pool sizes in vegetation and soil in five coniferous forest stands at the NITREX sites: Gardsjon (GD), Sweden, Klosterhede (KH), Denmark, Aber (AB), Wales, UK, Speuld (SP), the Netherlands, and Ysselsteyn (YS), the Netherlands. The sites span a N- deposition gradient from 13 to 59 kg N ha−1 yr−1. Measurements of soil N transformation rates by laboratory and field incubations were part of the site comparison. Further, results from 4–5 yr of NH4NO3 addition (35 kg N ha−1 yr−1) at low deposition sites (GD, KH, AB) and 6 yr of N removal (roofs) at high deposition sites (SP, YS) were included in the analysis. Significant correlations were found between a range of variables including N concentrations in foliage and litter, soil N transformation rates and forest floor characteristics. Using the methods from principal component analysis (PCA) these variables were summarized to an index of site N status that assigned the lowest N status to GD and the highest to YS. Site N status increased with N deposition with the exception that AB was naturally rich in N. Nitrate leaching was significantly correlated with N status but not correlated with N deposition. Forest floor mass and root biomass decreased with increased N status. Characteristics of the mineral soil were not correlated with vegetation and forest floor variables. High C N ratios in the mineral soil at the high-N deposition sites (SP, YS) suggest that the mineral soil pool changes slowly and need not change for N saturation to occur. Nitrogen transformation rates measured in laboratory incubations did not agree well with rates measured in the field except for a good correlation between ‘gross’ mineralization in the laboratory and ‘net’ mineralization in the field. The changes in N concentrations and fluxes after manipulation of N input followed the direction expected from the site comparison: increases at N addition and decreases at N removal sites. Nitrate leaching responded within the first year of treatment at all sites, whereas responses in vegetation and soil were delayed. Changes in N status by the manipulation treatments were small compared to the differences between sites. Changes in nitrate leaching were small at the low-N status sites and substantial at the high-N status sites. Nitrogen-limited and N-saturated forest ecosystems could be characterized quantitatively.

The Response of Soil Processes to Climate Change: Results from Manipulation Studies of Shrublands Across an Environmental Gradient

Predicted changes in climate may affect key soil processes such as respiration and net nitrogen (N) mineralization and thus key ecosystem functions such as carbon (C) storage and nutrient availability. To identify the sensitivity of shrubland soils to predicted climate changes, we have carried out experimental manipulations involving ecosystem warming and prolonged summer drought in ericaceous shrublands across a European climate gradient. We used retractable covers to create artificial nighttime warming and prolonged summer drought to 20-m2 experimental plots. Combining the data from across the environmental gradient with the results from the manipulation experiments provides evidence for strong climate controls on soil respiration, net N mineralization and nitrification, and litter decomposition. Trends of 0%–19% increases of soil respiration in response to warming and decreases of 3%–29% in response to drought were observed. Across the environmental gradient and below soil temperatures of 20°C at a depth of 5–10 cm, a mean Q10 of 4.1 in respiration rates was observed although this varied from 2.4 to 7.0 between sites. Highest Q10 values were observed in Spain and the UK and were therefore not correlated with soil temperature. A trend of increased accumulated surface litter mass loss was observed with experimental warming (2%– 22%) but there was no consistent response to experimental drought. In contrast to soil respiration and decomposition, variability in net N mineralization was best explained by soil moisture rather than temperature. When water was neither limiting or in excess, a Q10 of 1.5 was observed for net N mineralization rates. These data suggest that key soil processes will be differentially affected by predicted changes in rainfall pattern and temperature and the net effect on ecosystem functioning will be difficult to predict without a greater understanding of the controls underlying the sensitivity of soils to climate variables.

Novel Approaches to Study Climate Change Effects on Terrestrial Ecosystems in the Field: Drought and Passive Nighttime Warming

This article describes new approaches for manipulation of temperature and water input in the field. Nighttime warming was created by reflection of infrared radiation. Automatically operated reflective curtains covered the vegetation at night to reduce heat loss to the atmosphere. This approach mimicked the way climate change, caused by increased cloudiness and increased greenhouse gas emissions, alters the heat balance of ecosystems. Drought conditions were created by automatically covering the vegetation with transparent curtains during rain events over a 2–5-month period. The experimental approach has been evaluated at four European sites across a climate gradient. All sites were dominated (more than 50%) by shrubs of the ericaceous family. Within each site, replicated 4-m × 5-m plots were established for control, warming, and drought treatments and the effect on climate variables recorded. Results over a two-year period indicate that the warming treatment was successful in achieving an increase of the minimum temperatures by 0.4–1.2°C in the air and soil. The drought treatment resulted in a soil moisture reduction of 33%–82% at the peak of the drought. The data presented demonstrate that the approach minimizes unintended artifacts with respect to water balance, moisture conditions, and light, while causing a small but significant reduction in wind speed by the curtains. Temperature measurements demonstrated that the edge effects associated with the treatments were small. Our method provides a valuable tool for investigating the effects of climate change in remote locations with minimal artifacts.

Nonintrusive Field Experiments Show Different Plant Responses to Warming and Drought Among Sites, Seasons, and Species in a North–South European Gradient

We used a novel, nonintrusive experimental system to examine plant responses to warming and drought across a climatic and geographical latitudinal gradient of shrubland ecosystems in four sites from northern to southern Europe (UK, Denmark, The Netherlands, and Spain). In the first two years of experimentation reported here, we measured plant cover and biomass by the pinpoint method, plant 14C uptake, stem and shoot growth, flowering, leaf chemical concentration, litterfall, and herbivory damage in the dominant plant species of each site. The two years of approximately 1°C experimental warming induced a 15% increase in total aboveground plant biomass growth in the UK site. Both direct and indirect effects of warming, such as longer growth season and increased nutrient availability, are likely to be particularly important in this and the other northern sites which tend to be temperature-limited. In the water-stressed southern site, there was no increase in total aboveground plant biomass growth as expected since warming increases water loss, and temperatures in those ecosystems are already close to the optimum for photosynthesis. The southern site presented instead the most negative response to the drought treatment consisting of a soil moisture reduction at the peak of the growing season ranging from 33% in the Spanish site to 82% in The Netherlands site. In the Spanish site there was a 14% decrease in total aboveground plant biomass growth relative to control. Flowering was decreased by drought (up to 24% in the UK and 40% in Spain). Warming and drought decreased litterfall in The Netherlands site (33% and 37%, respectively) but did not affect it in the Spanish site. The tissue P concentrations generally decreased and the N/P ratio increased with warming and drought except in the UK site, indicating a progressive importance of P limitation as a consequence of warming and drought. The magnitude of the response to warming and drought was thus very sensitive to differences among sites (cold-wet northern sites were more sensitive to warming and the warm-dry southern site was more sensitive to drought), seasons (plant processes were more sensitive to warming during the winter than during the summer), and species. As a result of these multiple plant responses, ecosystem and community level consequences may be expected.

Natural abundance of 15N in forests across a nitrogen deposition gradient

Chronic atmospheric nitrogen deposition can alter the rate of internal nitrogen cycling and increase the magnitude of N leaching losses in forested ecosystems. As fractionation of nitrogen in favour of the lighter 14N occurs during various transformations associated with N-enrichment and nitrogen loss, it has been proposed that the 15N signal of vegetation may provide a useful tool in evaluating the past and current N status of forested ecosystems. A series of coniferous forests across a European nitrogen deposition gradient within the NITREX project provided an opportunity to test the relationships between nitrogen supply from atmospheric deposition and the relative 15N-enrichment of vegetation to soil, across a large geographical area. Most δ15N values for above- and below-ground tree components, soil at four depths, bulk precipitation and/or throughfall water and soil solution or outflow water values were within those observed elsewhere except for a few notable exceptions. There was a significant positive relationship between the δ15N enrichment of the tree foliage relative to the soil horizons (or the enrichment factor), and nitrogen flux in the throughfall if Aber forest, N. Wales, was excluded from the regression analysis. An unusually high enrichment factor at the Aber site indicated that a the high rate of N cycling at the site was in excess of that predicted from current N deposition. This was attributed to the effect of ploughing and tree planting on the relatively N- and clay-rich mineral horizons at Aber compared to other sites. Highly significant relationships (P < 0.01) between enrichment factors and parameters describing internal rates of N cycling, such as litterfall N flux and nitrification rates in upper soil horizons, supported this conclusion. There appears to be a strong link between the rate of N cycling and the δ15N enrichment factor, rather than N deposition or nitrate leaching per se. These results confirm the potential use of the δ15N enrichment factor to identify sites influenced by nitrogen deposition. However, consideration should be taken of other site characteristics and land management practises which also influence soil N dynamics and N cycling.

Predicting the Effects of Atmospheric Nitrogen Deposition in Conifer Stands: Evidence from the NITREX Ecosystem-Scale Experiments

ABSTRACT The NITREX project, which encompasses seven ecosystem-scale experiments in coniferous forests at the plot or catchment level in northwestern Europe, investigates the effect of atmospheric nitrogen (N) deposition in coniferous forests. The common factor in all of the experiments is the experimentally controlled change in N input over a period of 4–5 years. Results indicate that the status and dynamics of the forest floor are key components in determining the response of forests to altered N inputs. An empirical relationship between the carbon–nitrogen (C/N) ratio of the forest floor and retention of incoming N provides a simply measured tool through which the likely timing and consequences of changes in atmospheric N deposition for fresh waters may be predicted. In the terrestrial ecosystem, a 50% increase in tree growth is observed following the experimental reduction of N and sulfur inputs in a highly N-saturated site, illustrating the damaging effects of acidifying pollutants to tree health in some locations. Few biotic responses to the experimental treatments were observed in other NITREX sites, but the rapid response of water quality to changes in N deposition, and the link to acidification in sensitive areas, highlight the need for N-emission controls, irrespective of the long-term effects on tree health. The observed changes in ecosystem function in response to the experimental treatments have been considered within the framework of the current critical-load approach and thus contribute to the formulation of environmental policy.

The fate of 15N-labelled nitrogen deposition in coniferous forest ecosystems

Abstract As part of four European ecosystem manipulation experiments in coniferous forests, field-scale 15N tracer experiments have been carried out. The experiments involved a year-long addition of 15NH4+ and/or 15NO3− to throughfall at experimental plots with different N inputs. The fate of this applied 15N in the important ecosystems pools (trees, ground vegetation, forest floor and mineral soil), as well as in drainage was measured. About 10–30% of added 15N was taken up by the trees and 10–15% was retained in the mineral soil. Both retention efficiencies were found to be constant with N input. The part of 15N retained in the organic layer was relatively high (20–45% of applied) at low N inputs (0–30 kg N ha−1 yr−1) but low (10–20%) at high N inputs (30–80 kg N ha−1 yr−1). An inverse relationship between N input and the loss of 15N in drainage was found: drainage losses increased as a function of N input. These results suggest that increased N inputs exceed the capacity of the microbial population to retain throughfall-N in the organic layer, with the result that N leaching increases.

Microbial carbon and nitrogen dynamics in coniferous forest floor material collected along a European nitrogen deposition gradient

Abstract Microbial carbon (C) and nitrogen (N) cycling was studied in forest floor material from five coniferous forests situated along a gradient of N deposition across northwestern Europe. Results from the European NITREX project, a consortium of eight field-scale manipulation experiments with N deposition, indicated that three of these sites could be considered as N-saturated, whereas the other two sites were N-limited. Compared to the N-saturated sites, microbial C and N cycling in N-limited sites was characterized by low gross NH 4 + transformation rates, low respiration rates, high microbial C efficiencies and high microbial C:N ratios. Microbial NO 3 − immobilization was not detected in any of the forest floors studied here.

The chemolithotrophic ammonium-oxidizing community in a nitrogen-saturated acid forest soil in relation to ph-dependent nitrifying activity

Abstract The relationship between pH and nitrification in different layers of a nitrogen-saturated acid Douglas fir forest soil was studied. Nitrification potentials (on basis of dry weight) of overground degrading needles, litter and fermentation layers were much higher than those of the humus and upper mineral layers. In all layers, nitrate production was probably due to chemolithotrophic bacteria as it was inhibited by acetylene. The litter layer contained relatively high numbers (105 g dry soil−1) of acid-sensitive ammonium-oxidizing bacteria, whereas the numbers in the other layers were just above or lower than the detection limit (103 g dry soil−1) of the MPN method used. Measurements of nitrate production in soil suspensions indicated that a pH increase (pH 6 vs pH 4) stimulated ammonium oxidation in the litter layer but not or to a lesser extent in the fermentation and humus layers. It is argued that both acid-sensitive and acid-tolerant ammonium-oxidizing bacteria are contributing to nitrification in the litter layer. In the fermentation and humus layers acid-tolerant or even acidophilic bacteria are thought to be responsible for ammonium oxidation.