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Purpose Phosphorus (P) is a limiting nutrient in many managed forests. To further understand the risks and benefits of biochars as sustainable P source in forest management, an improved mechanistic understanding of its interactions in root systems is required. Methods A rhizobox experiment was conducted to observe root response of P. sylvestris (Scots pine) seedlings to different biochars in comparison to triple superphosphate (TSP) fertiliser as a P source. Three types of wood-derived biochar were compared: biochar from mixed softwood pellets (“Reference biochar”); from the vascular cambium zone of Picea sitchensis (“VCZ biochar”) and from mixed softwood pellets infused with TSP (“Processed biochar”). These biochars presented a range of available P from low to high. Seedling root development was spatially analysed using GIS software. Results The total length of P. sylvestris roots did not significantly differ between treatments. However, seedling roots showed strong preference for soil proximal to VCZ biochar and strong avoidance to TSP fertiliser. There was a milder avoidance effect for Processed biochar. Differences in root responses could be explained by available P: roots favored a moderate, sustained P source and avoided high available P sources. The avoidance effect can be attributed partially to lower soil pH around TSP fertiliser. Conclusion The extent concentration and duration of P availability affects the root response of P. sylverstris to P sources. Under P-deficient conditions, P. sylvestris root growth was markedly improved by introducing biochar with a certain P concentration, and VCZ biochar has potential as an effective source of P in forest establishment.

The stable nitrogen isotope (15 N) analysis of ammonium (NH4+) and nitrate (NO3–) is widely used in ecological research, providing insights into N cycling and its underlying regulating mechanisms in both aquatic and terrestrial ecosystems. To date, a large number of methods have been developed for the preparation and measurement of 15 N abundance of NH4+ and NO3– in liquid environmental samples at either natural abundance or enriched levels. However, these methods are all subject to certain specific advantages and limitations, and ecologists might be looking for an efficient way to select the most suitable methods in face of shifting sampling and analytical conditions. Based on our extensive review of these 15 N analysis methods we developed a decision support tool (DST) to provide quick and proper guidance for environmental researchers in finding the optimal method for preparing their liquid samples for 15 N analysis in NH4+ or NO3–. The DST is a decision tree based on several key criteria that users need to take into account when choosing the preferred sample preparation method for their samples. The criteria concern: the sample matrix, the 15 N abundance and the concentration of the target N species, the contamination by other N-containing chemicals, the isotopic fractionation, the availability of equipment, concerns about toxicity of reagents, and the preparation time. This work links field-scale experiments and laboratory 15 N analysis. Potential applications of our decision trees include 15 N studies ranging from natural abundance to tracer level in a wide range of terrestrial, freshwater and marine ecosystems.

Deforestation has a wide range of effects on hydrological and geochemical processes. Dissolved organic carbon (DOC) dynamics, a sensitive environmental change indicator, is expected to be affected by deforestation, with changes in atmospheric sulfur (S) deposition compounding this. However, how precisely anthropogenic disturbance (deforestation) under a declining atmospheric S input scenario affects the underlying spatiotemporal dynamics and relationships of river DOC and sulfate with hydro-climatological variables e.g., stream water temperature, runoff, pH, total dissolved iron (Fetot), and calcium (Ca2+) remains unclear. We, therefore, examined this issue within the TERENO Wüstebach catchment (Eifel, Germany), where partial deforestation had taken place in 2013. Wavelet transform coherence (WTC) analysis was applied based on a 10-year time series (2010–2020) from three sampling stations, whose (sub) catchment areas have different proportions of deforested area (W10: 31%, W14: 25%, W17: 3%). We found that water temperature and DOC, sulfate, and Fetot concentrations showed distinct seasonal patterns, with DOC averaging concentrations ranging from 2.23 (W17) to 4.56 (W10) mg L-1 and sulfate concentration ranging from 8.04 (W10) to 10.58 (W17) mg L-1. After clear-cut, DOC significantly increased by 59, 58% in the mainstream (W10, W14), but only 26% in the reference stream. WTC results indicated that DOC was negatively correlated with runoff and sulfate, but positively correlated with temperature, Ca2+, and Fetot. The negative correlation between DOC with runoff and sulfate was apparent over the whole examined 10-year period in W17 but did end in W10 and W14 after the deforestation. Sulfate was highly correlated with stream water temperature, runoff, and Fetot in W10 and W14 and with a longer lag time than W17. Additionally, pH was stronger correlated (higher R2) with sulfate and DOC in W17 than in W10 and W14. In conclusion, WTC analysis indicates that within this low mountainous forest catchment deforestation levels over 25% (W10 and W14) affected the coupling of S and C cycling substantially more strongly than “natural” environmental changes as observed in W17.

The capacity to forecast the effects of climate change on biodiversity largely relies on identifying traits capturing mechanistic relationships with the environment through standardized field experiments distributed across relevant spatial scales. The effects of short-term experimental manipulations on local communities may overlap with regional climate gradients that have been operating during longer time periods. However, to the best of our knowledge, there are no studies simultaneously assessing such long-term macroecological drivers with local climate manipulations. We analysed this issue with springtails (Class Collembola), one of the dominant soil fauna groups, in a standardized climate manipulation experiment conducted across six European countries encompassing broad climate gradients. We combined community data (near 20K specimens classified into 102 species) with 22 eco-morphological traits and reconstructed their phylogenetic relationships to track the evolution of adaptations to live at different soil depths, which is key to cope with desiccation. We then applied joint species distribution models to investigate the combined effect of the regional aridity gradient with the local experimental treatment (drought and warming) over the assembly of springtail communities and tested for significant trait–environment relationships mediating their community-level responses. Our results show (1) a convergent evolution in all three major collembolan lineages of species adapted to inhabit at different soil strata; (2) a clear signature of aridity selecting traits of more epigeic species at a biogeographical scale and (3) the association of short-term experimental drought with traits related to more euedaphic life-forms. The hemiedaphic condition would be the plesiomorphic state for Collembola while the adaptations for an epigeic life would have been secondarily gained. Epigeic springtails are not only more resistant to drought, but also have a higher dispersal capacity that allows them to seek more favourable micro-habitats after experiencing drier conditions. The observed relative edaphization of the springtail communities after short-term experimental drought may thus be a transient community response. The disparity between macroecological trends and fast community-level responses after climate manipulations highlights the need of simultaneously assessing long-term and short-term drivers at broad spatial scales to adequately interpret trait–environment relationships and better forecast biodiversity responses to climate change. Read the free Plain Language Summary for this article on the Journal blog.

Climate change is associated with an increased risk in the occurrence of wildfires. Forests store large amounts of carbon (C), which are threatened by these wildfires. Pyrogenic material produced after a wildfire constitutes an important part of the soil organic carbon pool in forest soils. Microorganisms play an important role in the cycling of C. This study investigated microbial activity in dissolved char from burned wood from two tree species in different stages of decay. The char from branches of beech and Norway spruce was produced under laboratory fire conditions and extracted in water after which microbial activity was measured for a 4-week period. Further stages of decay resulted in increased flammability with higher peak temperatures and combustion completeness. For the beech samples, further decay also resulted in a decrease of extractable C but a higher proportion of stable C. Further decay resulted in less respiration for beech and more for Norway spruce. With less C being respired, this points towards an increased C sequestration potential in the form of microbial C and microbial derived products for beech compared to Norway spruce. This study provided a workflow to assess the effects of dissolved char on microbial activity by mimicking natural fire conditions. It also indicated the need for future research to further elucidate the underlying mechanisms explaining why fire-originated dissolved char of wood in different decay stages influences microbial respiration with diverging effects per species.

High nitrogen (N) deposition levels, currently present in many industrial and agricultural regions of the world, can strongly affect the functioning of forest ecosystems. In a pine forest with strong N leaching, located in the Netherlands, we studied the long-term fate of a year-long NH4+ deposition cohort labeled with 15N. A high ambient and a low N deposition treatment had been established at the site by means of a roof and sprinklers. Resampling the N pools 19 years after labeling and 11 years after the last sampling, we found similar 15N deltas in needles, twigs and the LF1 organic soil layer of each treatment, indicating intensive N cycling among these pools. In the last 11 years, label recovery decreased in these labile pools, while recovery remained constant in wood and increased in bark. Together these aboveground vegetation pools retained less than 3% of the labeled N. In the organic layers, label recovery after 19 years decreased to 23% in both treatments, while in the mineral soil it increased from 4% to 13% (high N) and from 3% to 29% (low N treatment). Within the mineral soil of the high N treatment the labeled N was mainly found in fine roots, while in the low N treatment most N was incorporated in the two soil density fractions, shifting to the high density fraction with depth. This suggests a low capacity of the mineral soil at high N deposition to incorporate N. After the labeled N had been lost substantially in previous years, especially in the first, its presence remained constant in the last 11 years at 38% (high N) and 54% (low N treatment). Apparently, even in this strongly N leaching ecosystem, N once incorporated, was retained well and did not affect the input-output fluxes of the system.

Researchers use both experiments and observations to study the impacts of climate change on ecosystems, but results from these contrasting approaches have not been systematically compared for droughts. Using a meta-analysis and accounting for potential confounding factors, we demonstrate that aboveground biomass responded only about half as much to experimentally imposed drought events as to natural droughts. Our findings indicate that experimental results may underestimate climate change impacts and highlight the need to integrate results across approaches. In a meta-analysis comparing experimental versus observational studies of aboveground biomass responses to drought in grasslands, the authors show that effect sizes in experiments are 53% weaker than in observational studies, suggesting that experiments are underestimating drought responses.

Global industrial sulphur (S) dioxide emissions between 1900 to 1980 led to excessive S deposition and associated soil acidification. However, since introducing effective mitigation strategies, industrial S emissions have been significantly reduced, with concurrent reductions in S deposition. This has resulted in S deficiency in many croplands which now require supplementary S applications via fertilisers. We examined if such past differential atmospheric S inputs (‘legacy’) influence organic (or inorganic) S dynamics in current agricultural soils. We used a 62-year chronosequence of the reclaimed agricultural field after brown-coal mining (Inden, Germany) to sample topsoil (0-30 cm) from seven sites (representing the years 1956, 1971, 1985, 1995, 2005, 2011, and 2018). The dynamics of sulphur transformation were determined by adding 35S labelled methionine (Met) at 6, 24 and 48 h in an incubation experiment. The 35S-Met and 35S-SO4derived from labelled Metwere determined by measuring CaCl2-extractable 35S with or without BaCl2, the difference between the total added 35S-Met and the CaCl2-extractable 35S was recognized as the 35S immobilised in the microbial biomass. Results showed that soil S concentrations declined in a curvilinear pattern over the full chronosequence, from 0.27 (in 1956) to 0.11 g S kg-1 soil (in 2018). In contrast, soil C peaked in 1995 at 16 g C kg-1 soil, with the lowest values in 1956 at 10 g C kg-1 soil. For the site recultivated in 1985, transformation and S dynamics obviously differed from others. Here, compared with other sites, the 35S-SO4(inorganic S) concentrations (as % of the total 35S-Met added) peaked at 12, 29, 38% respectively, and 35S-Met (organic S) was the lowest at 35, 23, and 16%, respectively (at sampling times, 6, 24, and 48 h). The microbial biomass immobilized 53% of 35S-Met added to the soils in less than 6 h, and gradually released it as 35S-SO4 as incubation time increased. We conclude that organic S transformation in the soils was driven by the C rather S content, possible through differences in microbial C biomass, As such the effect of the S legacy in the soils could not be confirmed.

Shrubland ecosystems are vulnerable and typical ecosystems across European countries, but now they are facing a range of threats and an uncertain future because of climate change. Within the INCREASE project, six shrubland ecosystems along a geographical and climatic gradient across Europe from Wales and Denmark in the North, over The Netherlands to Hungary and Italy in the South, were exposed to ecosystem-level warming and extended periods of drought using automated curtain technologies. Sites ranged naturally from xeric to hydric. During our measurement period, mean annual precipitation (MAP) was reduced by 8-25%, while mean annual air temperature (MAT) was increased year-round by 0.2 - 0.9 ℃.Previously, Reinsch et al. (2017) * showed that aboveground net primary production (ANPP) was relatively resilient to the climate treatments while soil respiration (Rs) was reduced in xeric to mesic sites but increased in the hydric site. However, quantification of the gross primary production (GPP) or ecosystem respiration (Reco) rates and their responsiveness to climate manipulations have not previously been published. We will here present data from the six shrubland sites along the European climate gradient of the responses of GPP and Reco to drought and warming expressed as annual relative change (%) from the untreated control along a Gaussen index (GI). Results are in contrast to the previously reported decrease in Rs responsiveness with increased aridity. For both Reco and GPP rates, our preliminary results indicate that the more arid sites have a stronger, negative effect of drought suggesting different response patterns of autotrophic and heterotrophic components of the ecosystems.*Reinsch, S., Koller, E., Sowerby, A. et al. Shrubland primary production and soil respiration diverge along European climate gradient. Sci Rep 7, 43952 (2017). https://doi.org/10.1038/srep43952

In order to reduce the plastic accumulation in the environment, biodegradable plastics are attracting interest in the plastics market. However, the low thermal stability of most amorphous biodegradable polymers limits their application. With the aim of combining high glass transition temperature (T

Altres ajuts: Ramon Areces Foundation. Grant Number BEVP27P01A3355 Soil fauna play a fundamental role on key ecosystem functions like organic matter decomposition, although how local assemblages are responding to climate change and whether these changes may have consequences to ecosystem functioning is less clear. Previous studies have revealed that a continued environmental stress may result in poorer communities by filtering out the most sensitive species. However, these experiments have rarely been applied to climate change factors combining multiyear and multisite standardized field treatments across climatically contrasting regions, which has limited drawing general conclusions. Moreover, other facets of biodiversity, such as functional and phylogenetic diversity, potentially more closely linked to ecosystem functioning, have been largely neglected. Here, we report that the abundance, species richness, phylogenetic diversity, and functional richness of springtails (Subclass Collembola), a major group of fungivores and detritivores, decreased within 4 years of experimental drought across six European shrublands. The loss of phylogenetic and functional richness was higher than expected by the loss of species richness, leading to communities of phylogenetically similar species sharing evolutionary conserved traits. Additionally, despite the great climatic differences among study sites, we found that taxonomic, phylogenetic, and functional richness of springtail communities alone were able to explain up to 30% of the variation in annual decomposition rates. Altogether, our results suggest that the forecasted reductions in precipitation associated with climate change may erode springtail communities and likely other drought-sensitive soil invertebrates, thereby retarding litter decomposition and nutrient cycling in ecosystems

In order to reduce the plastic accumulation in the environment, biodegradable plastics are attracting interest in the plastics market. However, the low thermal stability of most amorphous biodegradable polymers limits their application. With the aim of combining high glass transition temperature (Tg), with good (marine) biodegradation a family of novel fully renewable poly(isosorbide-co-diol) oxalate (PISOX-diol) copolyesters was recently developed. In this study, the biodegradability of a representative copolyester, poly(isosorbide-co-1,6-hexanediol) oxalate (PISOX-HDO), with 75/25 mol ratio IS/HDO was evaluated at ambient temperature (25 °C) in soil and marine environment by using a Respicond system with 95 parallel reactors, based on the principle of frequently monitoring CO2 evolution. During 50 days incubation in soil and seawater, PISOX-HDO mineralised faster than cellulose. The ready biodegradability of PISOX-HDO is related to the relatively fast non-enzymatic hydrolysis of polyoxalates. To study the underlying mechanism of PISOX-HDO biodegradation, the non-enzymatic hydrolysis of PISOX-HDO and the biodegradation of the monomers in soil were also investigated. Complete hydrolysis was obtained in approximately 120 days (tracking the formation of hydrolysis products via 1H NMR). It was also shown that (enzymatic) hydrolysis to the constituting monomers is the rate-determining step in this biodegradation mechanism. These monomers can subsequently be consumed and mineralised by (micro)organisms in the environment much faster than the polyesters. The combination of high Tg (>100 °C) and fast biodegradability is quite unique and makes this PISOX-HDO copolyester ideal for short term applications that demand strong mechanical and physical properties.

High nitrogen (N) deposition levels, currently present in many industrial and agricultural regions of the world, can strongly affect the functioning of forest ecosystems. In a pine forest with strong N leaching, located in the Netherlands, we studied the long-term fate of a year-long NH

Increased anthropogenic nitrogen (N) inputs can alter the N cycle and affect forest ecosystem functions. The impact of increased N deposition depends among others on the ultimate fate of N in plant and soil N pools. Short-term studies (3–18 months) have shown that the organic soil layer was the dominant sink for N. However, longer time scales are needed to investigate the long-term fate of N. Therefore, the soils of four experimental forest sites across Europe were re-sampled ~ 2 decades after labelling with 15N. The sites covered a wide range of ambient N deposition varying from 13 to 58 kg N ha−1 year−1. To investigate the effects of different N loads on 15N recovery, ambient N levels were experimentally increased or decreased. We hypothesized that: (1) the mineral soil would become the dominant 15N sink after 2 decades, (2) long-term increased N deposition would lead to lower 15N recovery levels in the soil and (3) variables related to C dynamics would have the largest impact on 15N recovery in the soil. The results show that large amounts of the added 15N remain in the soil after 2 decades and at 2 out of 4 sites the 15N recovery levels are higher in the mineral soil than in the organic soil. The results show no clear responses of the isotopic signature to the changes in N deposition. Several environmental drivers are identified as controlling factors for long-term 15N recovery. Most drivers that significantly contribute to 15N recovery are strongly related to the soil organic matter (SOM) content. These findings are consistent with the idea that much of the added 15N is immobilized in the SOM. In the organic soil layer, we identify C stock, thickness of the organic layer, N-status and mean annual temperature of the forest sites as most important controlling factors. In the mineral soil we identify C stock, C content, pH, moisture content, bulk density, temperature, precipitation and forest stand age as most important controlling factors. Overall, our results show that these temperate forests are capable of retaining long-term increased N inputs preferably when SOM availability is high and SOM turnover and N availability are low.

Biodegradable plastics made from biopolymers (made in nature) or from bio-based polymers (made in a factory) are becoming increasingly important in replacing the massive amounts of conventional, non-degradable fossil-based plastics that have been produced and disposed over the past decades. In this review we compare the biodegradation rates and mechanisms of the bioplastics thermoplastic starch, cellulose acetate and lignin based bioplastics with the biodegradation rates and mechanisms of starch, cellulose and lignin, which are the unmodified biopolymers from which these bioplastics are produced. With this comparison we aim to determine to what extent the extensive knowledge on unmodified biopolymer biodegradation can be applied to the biodegradation of bioplastics (modified biopolymers) in the terrestrial environment. This knowledge is important, since it can be of great help in giving direction to the future research and development of bioplastics and for the development of bioplastic waste assessments and policies. We found that the similarities and differences in biodegradation are dependent on the structural changes imposed on a biopolymer during the bioplastic production process. A change in higher level structure, as found in thermoplastic starch, only resulted in a limited number of differences in the biodegradation process. However, when the chemical structure of a polymer is changed, as for cellulose acetate, different microorganisms and enzymes are involved in the biodegradation. Based on the cellulose acetate biodegradation process, a conceptual model was proposed that can be used as a starting point in predicting biodegradation rates of other chemically modified biopolymers used as bioplastics. Future bioplastic biodegradation research should focus on conducting long-term field experiments, since most studies are conducted in a laboratory setting and do not capture all processes occurring in the field situation. This applies even more to lignin based bioplastics, since very little experimental data were available on modified lignin biopolymer biodegradation.

We investigated the transfer of 15N into the soil via 15N uptake and release by tree roots, which involves the principles of the split-root technique. One half of the root system received an injection of (15NH4)2SO4 and the other half equivalent amounts of (NH4)2SO4 at 15N natural abundance level. 15N was transferred from one side of the root system (15N side) to the other side (14N side) and released into the soil. The method was conducted with Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies [L.] Karst). Two concentration levels of (NH4)2SO4 were used, corresponding with annual N deposition in the Netherlands (30 kg N ha–1) and a twelfth of that (2.5 kg N ha−1). Samples were taken 3 and 6 weeks after labelling and divided into needles + stem, roots, rhizosphere and bulk soil. Already 3 weeks after labelling, Scots pine took up 23.7 % of the low and 9.1 % of the high amounts of 15N, while Norway spruce took up 21.5 and 32.1 %, respectively. Both species transported proportions of 15N to the rhizosphere (0.1–0.2 %) and bulk soil (0.3–0.9 %). The method is a useful tool to investigate the fate of root-derived N in soils, for example, for the formation of stable forms of soil organic matter.

Climate change will alter precipitation patterns with consequences for soil C cycling. An understanding of how fluctuating soil moisture affects microbial processes is therefore critical to predict responses to future global change. We investigated how long-term experimental field drought influences microbial tolerance to lower moisture levels (“resistance”) and ability to recover when rewetted after drought (“resilience”), using soils from a heathland which had been subjected to experimental precipitation reduction during the summer for 18 years. We tested whether drought could induce increased resistance, resilience, and changes in the balance between respiration and bacterial growth during perturbation events, by following a two-tiered approach. We first evaluated the effects of the long-term summer drought on microbial community functioning to drought and drying–rewetting (D/RW), and second tested the ability to alter resistance and resilience through additional perturbation cycles. A history of summer drought in the field selected for increased resilience but not resistance, suggesting that rewetting after drought, rather than low moisture levels during drought, was the selective pressure shaping the microbial community functions. Laboratory D/RW cycles also selected for communities with a higher resilience rather than increased resistance. The ratio of respiration to bacterial growth during D/RW perturbation was lower for the field drought-exposed communities and decreased for both field treatments during the D/RW cycles. This suggests that cycles of D/RW also structure microbial communities to respond quickly and efficiently to rewetting after drought. Our findings imply that microbial communities can adapt to changing climatic conditions and that this might slow the rate of soil C loss predicted to be induced by future cyclic drought. (Less)

Well defined productivity-precipitation relationships of ecosystems are needed as benchmarks for the validation of land-models used for future projections. The productivity-precipitation relationship may be studied in two ways: the spatial approach relates differences in productivity to those in precipitation among sites along a precipitation gradient (the spatial fit, with a steeper slope); the temporal approach relates inter-annual productivity changes to variation in precipitation within sites (the temporal fits, with flatter slopes). Precipitation-reduction experiments in natural ecosystems represent a complement to the fits, because they can reduce precipitation below the natural range and are thus well suited to study potential effects of climate drying. Here, we analyze the effects of dry treatments in eleven multi-year precipitation-manipulation experiments, focusing on changes in the temporal fit. We expected that structural changes in the dry treatments would occur in some experiments, thereby reducing the intercept of the temporal fit and displacing the productivity-precipitation relationship downward the spatial fit. The majority of experiments (72%) showed that dry treatments did not alter the temporal fit. This implies that current temporal fits are to be preferred over the spatial fit to benchmark land-model projections of productivity under future climate within the precipitation ranges covered by the experiments. Moreover, in two experiments, the intercept of the temporal fit unexpectedly increased due to mechanisms that reduced either water- or nutrient losses. The expected decrease of the intercept was observed in only one experiment, and only when distinguishing between the late and the early phases of the experiment. This implies that we currently do not know at which precipitation-reduction level or at which experimental duration structural changes will start to alter ecosystem productivity. Our study highlights the need for experiments with multiple, including more extreme, dry treatments, to identify the precipitation boundaries within which the current temporal fits remain valid.

Worldwide, forests have absorbed around 30% of global anthropogenic emissions of carbon dioxide (CO2) annually, thereby acting as important carbon (C) sinks. It is proposed that leaving large fragments of dead wood, coarse woody debris (CWD), in forest ecosystems may contribute to the forest C sink strength. CWD may take years to centuries to degrade completely, and non-respired C from CWD may enter the forest soil directly or in the form of dissolved organic C. Although aboveground decomposition of CWD has been studied frequently, little is known about the relative size, composition and fate of different C fluxes from CWD to soils under various substrate-specific and environmental conditions. Thus, the exact contribution of C from CWD to C sequestration within forest soils is poorly understood and quantified, although understanding CWD degradation and stabilization processes is essential for effective forest C sink management. This review aims at providing insight into these processes on the interface of forest ecology and soil science, and identifies knowledge gaps that are critical to our understanding of the effects of CWD on the forest soil C sink. It may be seen as a “call-to-action” crossing disciplinary boundaries, which proposes the use of compound-specific analytical studies and manipulation studies to elucidate C fluxes from CWD. Carbon fluxes from decaying CWD can vary considerably due to interspecific and intraspecific differences in composition and different environmental conditions. These variations in C fluxes need to be studied in detail and related to recent advances in soil C sequestration research. Outcomes of this review show that the presence of CWD may enhance the abundance and diversity of the microbial community and constitute additional fluxes of C into the mineral soil by augmented leaching of dissolved organic carbon (DOC). Leached DOC and residues from organic matter (OM) from later decay stages have been shown to be relatively enriched in complex and microbial-derived compounds, which may also be true for CWD-derived OM. Emerging knowledge on soil C stabilization indicates that such complex compounds may be sorbed preferentially to the mineral soil. Moreover, increased abundance and diversity of decomposer organisms may increase the amount of substrate C being diverted into microbial biomass, which may contribute to stable C pools in the forest soil.

A catchment-scale mass-balance model of linked carbon and nitrogen cycling in ecosystems has been developed for simulating leaching losses of inorganic nitrogen. The model (MERLIN) considers linked biotic and abiotic processes affecting the cycling and storage of nitrogen. The model is aggregated in space and time and contains compartments intended to be observable and/or interpretable at the plot or catchment scale. The structure of the model includes the inorganic soil, a plant compartment and two soil organic compartments. Fluxes in and out of the ecosystem and between compartments are regulated by atmospheric deposition, hydrological discharge, plant uptake, litter production, wood production, microbial immobilization, mineralization, nitrification, and denitrification. Nitrogen fluxes are controlled by carbon productivity, the C:N ratios of organic compartments and inorganic nitrogen in soil solution. Inputs required are: 1) temporal sequences of carbon fluxes and pools- 2) time series of hydrological discharge through the soils, 3) historical and current external sources of inorganic nitrogen; 4) current amounts of nitrogen in the plant and soil organic compartments; 5) constants specifying the nitrogen uptake and immobilization characteristics of the plant and soil organic compartments; and 6) soil characteristics such as depth, porosity, bulk density, and anion/cation exchange constants. Outputs include: 1) concentrations and fluxes of NO3 and NH4 in soil solution and runoff; 2) total nitrogen contents of the organic and inorganic compartments; 3) C:N ratios of the aggregated plant and soil organic compartments; and 4) rates of nitrogen uptake and immobilization and nitrogen mineralization. The behaviour of the model is assessed for a combination of land-use change and nitrogen deposition scenarios in a series of speculative simulations. The results of the simulations are in broad agreement with observed and hypothesized behaviour of nitrogen dynamics in growing forests receiving nitrogen deposition.

Monomeric organic nitrogen (N) compounds such as free amino acids (FAAs) are an important resource for both plants and soil microorganisms and a source of ammonium (NH4+) via microbial FAA mineralization. We compared gross FAA dynamics with gross N mineralization in a Dutch heathland soil using a 15N tracing technique. A special focus was made on the effects of climate change factors warming and drought, followed by rewetting. Our aims were to (1) compare FAA mineralization (NH4+ production from FAAs) with gross N mineralization, (2) assess gross FAA production rate (depolymerization) and turnover time relative to gross N mineralization rate, and (3) assess the effects of a 14 years of warming and drought treatment on these rates. The turnover of FAA in the soil was ca. 3 h, which is almost 2 orders of magnitude faster than that of NH4+ (i.e. ca. 4 days). This suggests that FAA is an extensively used resource by soil microorganisms. In control soil (i.e. no climatic treatment), the gross N mineralization rate (10 ± 2.9 μg N g−1 day−1) was 8 times smaller than the total gross FAA production rate of five AAs (alanine, valine, leucine, isoleucine, proline: 127.4 to 25.0 μg N g−1 day−1). Gross FAA mineralization (3.4 ± 0.2 μg N g−1 day−1) contributed 34% to the gross N mineralization rate and is therefore an important component of N mineralization. In the drought treatment, a 6–29% reduction in annual precipitation caused a decrease of gross FAA production by 65% and of gross FAA mineralization by 41% compared to control. On the other hand, gross N mineralization was unaffected by drought, indicating an increased mineralization of other soil organic nitrogen (SON) components. A 0.5–1.5 °C warming did not significantly affect N transformations, even though gross FAA production declined. Overall our results suggest that in heathland soil exposed to droughts a different type of SON pool is mineralized. Furthermore, compared to agricultural soils, FAA mineralization was relatively less important in the investigated heathland. This indicates more complex mineralization dynamics in semi-natural ecosystems.

We investigated the transfer of

imbalance p paper contact with: Marc Estiarte, m.estiarte@creaf.uab.cat Above- and belowground carbon (C) stores of terrestrial ecosystems are vulnerable to environmental change. Ecosystem C balances in response to environmental changes have been quantified at individual sites, but the magnitudes and directions of these responses along environmental gradients remain uncertain. Here we show the responses of ecosystem C to 8-12 years of experimental drought and night-time warming across an aridity gradient spanning seven European shrublands using indices of C assimilation (aboveground net primary production: aNPP) and soil C efflux (soil respiration: Rs). The changes of aNPP and Rs in response to drought indicated that wet systems had an overall risk of increased loss of C but drier systems did not. Warming had no consistent effect on aNPP across the climate gradient, but suppressed Rs more at the drier sites. Our findings suggest that above- and belowground C fluxes can decouple, and provide no evidence of acclimation to environmental change at a decadal timescale. aNPP and Rs especially differed in their sensitivity to drought and warming, with belowground processes being more sensitive to environmental change.

As a key component of the carbon cycle, soil CO2 efflux (SCE) is being increasingly studied to improve our mechanistic understanding of this important carbon flux. Predicting ecosystem responses to climate change often depends on extrapolation of current relationships between ecosystem processes and their climatic drivers to conditions not yet experienced by the ecosystem. This raises the question of to what extent these relationships remain unaltered beyond the current climatic window for which observations are available to constrain the relationships. Here, we evaluate whether current responses of SCE to fluctuations in soil temperature and soil water content can be used to predict SCE under altered rainfall patterns. Of the 58 experiments for which we gathered SCE data, 20 were discarded because either too few data were available or inconsistencies precluded their incorporation in the analyses. The 38 remaining experiments were used to test the hypothesis that a model parameterized with data from the control plots (using soil temperature and water content as predictor variables) could adequately predict SCE measured in the manipulated treatment. Only for 7 of these 38 experiments was this hypothesis rejected. Importantly, these were the experiments with the most reliable data sets, i.e., those providing high-frequency measurements of SCE. Regression tree analysis demonstrated that our hypothesis could be rejected only for experiments with measurement intervals of less than 11 days, and was not rejected for any of the 24 experiments with larger measurement intervals. This highlights the importance of high-frequency measurements when studying effects of altered precipitation on SCE, probably because infrequent measurement schemes have insufficient capacity to detect shifts in the climate dependencies of SCE. Hence, the most justified answer to the question of whether current moisture responses of SCE can be extrapolated to predict SCE under altered precipitation regimes is "no" – as based on the most reliable data sets available. We strongly recommend that future experiments focus more strongly on establishing response functions across a broader range of precipitation regimes and soil moisture conditions. Such experiments should make accurate measurements of water availability, should conduct high-frequency SCE measurements, and should consider both instantaneous responses and the potential legacy effects of climate extremes. This is important, because with the novel approach presented here, we demonstrated that, at least for some ecosystems, current moisture responses could not be extrapolated to predict SCE under altered rainfall conditions.

Soil respiration (SR) is a major component of the global carbon cycle and plays a fundamental role in ecosystem feedback to climate change. Empirical modelling is an essential tool for predicting ecosystem responses to environmental change, and also provides important data for calibrating and corroborating process-based models. In this study, we evaluated the performance of three empirical temperature-SR response functions (exponential, Lloyd-Taylor and Gaussian) at seven shrublands located within three climatic regions (Atlantic, Mediterranean and Continental) across Europe. We investigated the performance of SR models by including the interaction between soil moisture and soil temperature. We found that the best fit for the temperature functions depended on the site-specific climatic conditions. Including soil moisture, we identified thresholds in the three different response functions that improved the model fit in all cases. The direct soil moisture effect on SR, however, was weak at the annual time scale. We conclude that the exponential soil temperature function may only be a good predictor for SR in a narrow temperature range, and that extrapolating predictions for future climate based on this function should be treated with caution as modelled outputs may underestimate SR. The addition of soil moisture thresholds improved the model fit at all sites, but had a far greater ecological significance in the wet Atlantic shrubland where a fundamental change in the soil CO2 efflux would likely have an impact on the whole carbon budget.

Heathlands are cultural landscapes which are managed through cyclical cutting, burning or grazing practices. Understanding the carbon (C) fluxes from these ecosystems provides information on the optimal management cycle time to maximise C uptake and minimise C output. The interpretation of field data into annual C loss values requires the use of soil respiration models. These generally include model variables related to the underlying drivers of soil respiration, such as soil temperature, soil moisture and plant activity. Very few studies have used selection procedures in which structurally different models are calibrated, then validated on separate observation datasets and the outcomes critically compared. We present thorough model selection procedures to determine soil heterotrophic (microbial) and autotrophic (root) respiration for a heathland chronosequence and show that soil respiration models are required to correct the effect of experimental design on soil temperature. Measures of photosynthesis, plant biomass, photosynthetically active radiation, root biomass, and microbial biomass did not significantly improve model fit when included with soil temperature. This contradicts many current studies in which these plant variables are used (but not often tested for parameter significance). We critically discuss a number of alternative ecosystem variables associated with soil respiration processes in order to inform future experimental planning and model variable selection at other heathland field sites. The best predictive model used a generalized linear multi-level model with soil temperature as the only variable. Total annual soil C loss from the young, middle and old communities was calculated to be 650, 462 and 435 g C m−2 yr−1, respectively.

Studies of biological responses in the terrestrial environment to rapid changes in climate have mostly been concerned with aboveground biota, whereas less is known of belowground organisms. The present study focuses on mites and springtails of heathland ecosystems and how the microarthropod community has responded to simulated climate change in a long-term field experiment. Increased temperature and repeated drought was applied for 13 years to field plots located in Wales, The Netherlands and Denmark representing sites of contrasting climatic conditions with respect to precipitation and temperature. This approach provided an opportunity to study biological responses on a local (within sites) and regional scale. Warming treatments increasing night time temperature (0.3-1 °C higher than ambient at 5 cm soil depth) had no detectable effects on the microarthropod communities. Increased intensity and frequency of drought had only weak persistent effects on springtail species composition, but practically no effect on major mite groups (Oribatida, Prostigmata or Mesostigmata) suggesting that ecosystem functions of microarthropods may only be transiently impacted by repeated spring or summer drought.

Field experiments that expose terrestrial ecosystems to climate change factors by manipulations are expensive to maintain, and typically only last a few years. Plant biomass is commonly used to assess responses to climate treatments and to predict climate change impacts. However, response to the treatments might be considerably different between the early years and a decade later. The aim of this data analysis was to develop and apply a method for evaluating changes in plant biomass responses through time, in order to provide a firm basis for discussing how the ‘short-term’ response might differ from the ‘long-term’ response. Across 22 sites situated in the northern hemisphere, which covered three continents, and multiple ecosystems (grasslands, shrublands, moorlands, forests, and deserts), we evaluated biomass datasets from long-term experiments with exposure to elevated CO2 (eCO2), warming, or drought. We developed methods for assessing biomass response patterns to the manipulations using polynomial and linear (piecewise) model analysis and linked the responses to site-specific variables such as temperature and rainfall. Polynomial patterns across sites indicated changes in response direction over time under eCO2, warming, and drought. In addition, five distinct pattern types were confirmed within sites: ‘no response’, ‘delayed response’, ‘directional response’, ‘dampening response’, and ‘altered response’ patterns. We found that biomass response direction was as likely to change over time as it was to be consistent, and therefore suggest that climate manipulation experiments should be carried out over timescales covering both short- and long-term responses, in order to realistically assess future impacts of climate change.

© 2016, National Academy of Sciences. All rights reserved. The respiratory release of carbon dioxide (CO2) from soil is a major yet poorly understood flux in the global carbon cycle. Climatic warming is hypothesized to increase rates of soil respiration, potentially fueling further increases in global temperatures. However, despite considerable scientific attention in recent decades, the overall response of soil respiration to anticipated climatic warming remains unclear. We synthesize the largest global dataset to date of soil respiration, moisture, and temperature measurements, totaling >3,800 observations representing 27 temperature manipulation studies, spanning nine biomes and over 2 decades of warming. Our analysis reveals no significant differences in the temperature sensitivity of soil respiration between control and warmed plots in all biomes, with the exception of deserts and boreal forests. Thus, our data provide limited evidence of acclimation of soil respiration to experimental warming in several major biome types, contrary to the results from multiple single-site studies. Moreover, across all nondesert biomes, respiration rates with and without experimental warming follow a Gaussian response, increasing with soil temperature up to a threshold of ∼25 °C, above which respiration rates decrease with further increases in temperature. This consistent decrease in temperature sensitivity at higher temperatures demonstrates that rising global temperatures may result in regionally variable responses in soil respiration, with colder climates being considerably more responsive to increased ambient temperatures compared with warmer regions. Our analysis adds a unique cross-biome perspective on the temperature response of soil respiration, information critical to improving our mechanistic understanding of how soil carbon dynamics change with climatic warming.

To study the long-term fate of deposited ammonium (NH4 +) in a Scots pine forest stand under high nitrogen (N) deposition in the Netherlands we re-sampled the plots of a 15N tracer experiment with high (i.e. ambient) and lowered N deposition in this stand 8 years after application of the tracer. The results were compared with results obtained 7 years earlier. In the 7 years between the samplings the 15N deltas of needles, twigs and upper organic soil layer had converged to similar values still above the natural 15N abundance, suggesting equilibration as a result of intensive cycling of N among these pools. Bark and wood had lower deltas than needles and twigs, but if the label found was attributed to tissue synthesized since the start of the labeling only, bark values were similar to needles and twigs, whereas wood values were higher indicating retranslocation of N into older wood. Mineral soil lost all 15N label it had accumulated after 1 year indicating that this label had not been strongly bound. The first year the low N treatment had retained more of the labeled NH4 + deposition than the high N treatment, but in the seven subsequent years relatively more label was retained in the latter. This better retention after 7 years was ascribed to a larger fraction of label taken up by the vegetation in the high N treatment. This shows that the vegetation can affect the label dynamics despite the fact that only a relatively small amount of label was present in the aboveground vegetation.

Studies of biological responses in the terrestrial environment to rapid changes in climate have mostly been concerned with above-ground biota, whereas less is known of belowground organisms. The present study focuses on enchytraeids (Oligochaeta) of heathland ecosystems and how the enchytraeid community has responded to simulated climate change in a long-term field experiment. Either increased temperature or repeated drought was applied for 13 years to field plots located in Wales, The Netherlands and Denmark representing a gradient in precipitation and annual temperature fluctuations thereby providing an opportunity to study biological responses on a local (within sites) and regional scale. Warming treatments increasing night-time temperature (0.5–1 °C higher than ambient at 5 cm soil depth) had no detectable effects on the enchytraeid communities. Increased intensity and frequency of drought had rather weak persistent effects on total enchytraeid abundance suggesting that ecosystem functions of enchytraeids may only be transiently impacted by repeated spring or summer drought. However, drought treatment had persistent negative effects on species richness and community structure across sites. Drought treated plots harboured only 35–65% of the species present in control plots, and the reduction of species richness was most pronounced at the driest sites. It is discussed that soil invertebrates, due to their weak migratory potential, may be more liable to extinction under changing climatic conditions than above-ground species, and therefore consequences of climate change to soil organisms need particular attention in future research.

Acid atmospheric emissions within Europe and North America have decreased strongly since 1985 and most recent acidification studies have focused on the changes occurring within ecosystems as a result of this decreased deposition. This current study documents a soil acidification trend under ambient N deposition conditions over a 13 year period, suggesting that acidification continues to be a process of concern at this Calluna vulgaris dominated heathland with an acidic sandy soil. The annual manipulation of climatic conditions on this heathland simulated the predicted summer rainfall reduction (drought) and resulted in a long term retardation of the soil acidification trend. The pH of the soil solution significantly decreased over the course of the trial for both treatments, however, in the final two years the decline continued only in the Control treatment. This retardation is primarily associated with the reduction in rainfall leading to lower drainage rates, reduced loss of cations and therefore reduced lowering of the soil acid neutralizing capacity (ANC). However, a change in the underlying mechanisms also indicated that N transformations became less important in the Drought treatment. This change corresponded to an increase in groundcover of an air-pollution tolerant moss species and it is hypothesized that this increasing moss cover filtered an increasing quantity of deposited N, thus reducing the N available for transformation. A soil acidification lag time is expected to increase between the two treatments due to the cumulative disparity in cation retention and rates of proton formation. To the authors knowledge, this is the first study in which such acidification trends have been demonstrated in a field-scale climate manipulation experiment.

Effects of anthropogenic nitrogen (N) deposition and the ability of terrestrial ecosystems to store carbon (C) depend in part on the amount of N retained in the system and its partitioning among plant and soil pools. We conducted a meta-analysis of studies at 48 sites across four continents that used enriched 15N isotope tracers in order to synthesize information about total ecosystem N retention (i.e., total ecosystem 15N recovery in plant and soil pools) across natural systems and N partitioning among ecosystem pools. The greatest recoveries of ecosystem 15N tracer occurred in shrublands (mean, 89.5%) and wetlands (84.8%) followed by forests (74.9%) and grasslands (51.8%). In the short term (< 1 week after 15N tracer application), total ecosystem 15N recovery was negatively correlated with fine-root and soil 15N natural abundance, and organic soil C and N concentration but was positively correlated with mean annual temperature and mineral soil C:N. In the longer term (3-18 months after 15N tracer application), total ecosystem 15N retention was negatively correlated with foliar natural-abundance 15N but was positively correlated with mineral soil C and N concentration and C:N, showing that plant and soil natural-abundance 15N and soil C:N are good indicators of total ecosystem N retention. Foliar N concentration was not significantly related to ecosystem 15N tracer recovery, suggesting that plant N status is not a good predictor of total ecosystem N retention. Because the largest ecosystem sinks for 15N tracer were below ground in forests, shrublands, and grasslands, we conclude that growth enhancement and potential for increased C storage in aboveground biomass from atmospheric N deposition is likely to be modest in these ecosystems. Total ecosystem 15N recovery decreased with N fertilization, with an apparent threshold fertilization rate of 46 kg N x ha(-1) x yr(-1) above which most ecosystems showed net losses of applied 15N tracer in response to N fertilizer addition.

The effects of wetting and drying and inorganic nitrogen (N) addition on carbon (C) and N turnover in fresh Douglas fir litter (Speuld forest, the Netherlands) were investigated. Litter was incubated for 9 days in the laboratory, receiving different moisture and N addition treatments. Following the additions, a series of reactions were observed of which most notable were a rapid retention of added ammonium and nitrate (NO3-) and a sudden increase in CO2 respiration. For the rewetted-and-moist incubations, respiration levels remained elevated, N was net immobilized and nitrous oxide (N2O) production increased throughout the experiment. About 80% of the NO3- produced was lost again as N2O. In the rewetted-and-dried incubations, respiration decreased during the drying phase; no clear patterns in N mineralization were detected; and N2O production remained at constant levels, but still resulted in gaseous loss for half of the NO3- net produced. The experiments thus revealed two important NO3- sinks in LF1 litter, namely rapid retention of added NO3- and gaseous loss as N2O. The maximum NO3- loss via these sinks was estimated at 2 kg-N ha-1 yr-1, which is small compared to annual NO3- leaching at 90 cm soil depth (31 kg-N ha-1 yr-1).

We investigated the carbon (C) mineralisation and nitrogen (N) dynamics in litter from a Corsican pine forest in response to individual and combined additions of aluminium (Al), condensed tannin (extracted from fresh Corsican pine needles) and hydrolysable tannin (commercial tannic acid). Production rates of CO 2 , NH 4 + and NO 3 − concentrations, tannin concentrations and Al speciation were determined at various time intervals during a 28-day incubation experiment. The addition of Al decreased CO 2 production and shut down nitrification. Exchangeable NH 4 + strongly increased in the Al-amended litter, likely due to (i) decreased microbial uptake of NH 4 + , (ii) the inhibition of nitrification and (iii) competition for soil organic matter (SOM) binding sites by Al. Both tannin species affected C mineralisation and/or N dynamics, be it in different ways. Addition of tannic acid led to a strong increase of the C mineralisation rate and microbial uptake of N, caused by rapid degradation of this labile tannin and subsequent increased microbial nutrient demand. Net immobilisation of N occurred as long as one week after addition. Condensed tannin was not consumed but probably strongly bound to (nitrogenous) SOM compounds, forming recalcitrant complexes and decreasing net N mineralisation. Complexation of Al by tannins in solution before addition to the litter mitigated the Al-induced release of exchangeable NH 4 + . In the case of condensed tannin with complexed Al, this was due to detoxification of Al through complexation. Increased microbial demand for N likely played a major role in decreased NH 4 + accumulation in the samples to which tannic acid with complexed Al was added. Nitrification was shut down despite of the complexation of Al by either condensed tannin or tannic acid.

Current predictions of climate change include altered rainfall patterns throughout Europe, continental USA and areas such as the Amazon. The effect of this on soil carbon efflux remains unclear although several modelling studies have highlighted the potential importance of drought for carbon storage. To test the importance of drought and more importantly, repeated drought year-on-year, we used automated retractable curtains to exclude rain and produce repeated summer drought in three heathlands at varying moisture conditions. This included a hydric system limited by water-excess (in the UK) and two mesic systems with seasonal water-limitation in Denmark (DK) and the Netherlands (NL). The experimental rainfall reductions were set to reflect single year droughts observed in the last decade with exclusion of rain for 2–3 months of the year resulting in a 20–26% reduction in annual rainfall and 23–38% reduction in mean soil moisture during the drought period. Unexpectedly, sustained reduction in soil moisture over the winter (between drought periods) was also observed at all three sites, along with a reduction in the maximum water holding capacity attained. Three hypotheses are discussed which may have contributed to this lack of recovery in soil moisture; hydrophobicity of soil organic matter, increased water use by plants and increased cracking of the soil. The responses of soil respiration to this change in soil moisture varied between the sites; decreased rates were observed at the water limited NL and DK sites whilst they increased at the UK site. Reduced sensitivity of soil respiration to soil temperature was observed at soil moisture contents above 55% at the UK site and below 20% and 13% at the NL and DK sites respectively. Soil respiration rates recovered to pre-drought levels in the NL and DK sites during the winter rewetting period that indicates any change in soil C storage due to changes in soil C efflux may be short-lived in these mesic systems. In contrast, in the hydric UK site after 2 years of drought treatment, the persistent reduction in soil moisture throughout the year, resulted in a year-round increase in soil respiration flux; a response which accelerated over time to 40% above control levels. These findings suggest that carbon-rich soils with high organic matter content may act as a significant source of CO2 to the atmosphere following repeated summer drought. Non-recovery of soil moisture and a persistent increase in soil respiration may be the primary mechanism underlying the reported substantial losses of soil carbon from UK organic soils over the last 20 years. These findings indicate that the water status of an ecosystem will be a critical factor to consider in determining the impact of drought on soil carbon fluxes and storage.

The development of anoxic conditions in forest litter and the relation with nitrous oxide (N2O) production and emission rates are not completely understood. Water content is an important factor in the regulation of N2O production due to its effect on the development of anoxic conditions. A combination of simulation modeling and incubation experiments was used to study (1) O2 concentrations in water and organic matter at various water saturation fractions of inter-particle pores in Douglas fir litter (F2 horizon), (2) the relationship between N2O production and moisture content of litter and (3) to test whether diffusion constraints of nitrate (NO3¿) could have explained measured N2O production rates within litter fragments. Model simulations showed that the occurrence of high N2O production rates in samples with extremely high water contents coincided with the development of anoxic conditions in water-filled inter-particle pores. Measured N2O production rates started to increase exponentially after 1¿2 days in glucose-amended samples, during which substantial microbial growth was established. For these latter samples model simulations showed that the increase in O2 consumption due to microbial growth lead to anoxic conditions in water-filled pores at locations which were far from the O2 saturated air-filled pores. It was concluded that anoxic conditions in water-filled pores was the crucial factor for the development of high N2O production rates. Diffusion limitation of NO3¿ and glucose were estimated to be negligible in the highly fragmented litter material used. The occurrence of diffusion limitation depended on litter particle size, the NO3¿ reduction potential and the NO3¿ concentration. Therefore, diffusion limitation together with N2O production in litter cannot be neglected under field conditions with a low NO3¿ concentration or a high NO3¿ reduction potential.

To evaluate the effect of climate change on ecosystem functioning, the temperature and moisture response of microbial C, N, and P transformations during decomposition of Calluna vulgaris (L.) Hull. litter was studied in a laboratory incubation experiment. The litter originated from a dry heathland in the Netherlands where P limited vegetation growth. Fresh litter was incubated at 5, 10, 15, or 20°C and at a moisture content of 50, 100, or 200% in a full factorial design. Microbial nutrient transformations and activity were evaluated during two successive periods: an initial period of 48 days characterized by microbial growth and a second period from 48 to 206 days in which microbial growth declined significantly. Temperature and moisture response of respiration rate, the metabolic quotient (qCO2), C, N, and P immobilization, net N and P mineralization and nitrification rates were evaluated by performing linear regressions. Microbial nutrient transformations and microbial activity depended both on temperature and moisture. In the first period, the respiration rate, qCO2, microbial C and N immobilization, net P mineralization, net N mineralization and net nitrification rates were more strongly affected by temperature, while the microbial P immobilization rate was more strongly affected by moisture. The respiration rate, qCO2, P immobilization rate, net P and N mineralization rate, and nitrification rate increased with temperature and moisture, while the C and N immobilization rate decreased with increasing temperature and increased with moisture. In the second period, C, N, and P immobilization and net N and P mineralization rates were significantly lower. The respiration rate and qCO2 continued to increase with temperature and moisture, but C and N immobilization rates increased with temperature and declined with increasing moisture. Net P mineralization rate decreased at higher temperature and moisture, and nitrification rate declined with increasing temperature and increased with moisture. It was concluded that plant growth in these P-limited systems is very sensitive to climate change as it strongly relies on the competition for P with microbes, and temperature and moisture have a large effect on the immobilization rate of available P.

Human domination of the biosphere includes changes to disturbance regimes, which push many ecosystems towards early-successional states. Ecological theory predicts that early-successional ecosystems are more sensitive to perturbations than mature systems, but little evidence supports this relationship for the perturbation of climate change. Here we show that vegetation (abundance, species richness and species composition) across seven European shrublands is quite resistant to moderate experimental warming and drought, and responsiveness is associated with the dynamic state of the ecosystem, with recently disturbed sites responding to treatments. Furthermore, most of these responses are not rapid (2-5 years) but emerge over a longer term (7-14 years). These results suggest that successional state influences the sensitivity of ecosystems to climate change, and that ecosystems recovering from disturbances may be sensitive to even modest climatic changes. A research bias towards undisturbed ecosystems might thus lead to an underestimation of the impacts of climate change.

Pine litter amended with either tannic acid (TA) or condensed tannins (CTs) was studied to assess the effects on C, N and P mineralization in relation to the fate of tannins by incubation experiments during various time intervals. TA induced a rapid short-term effect resulting in high C respiration and net N and P immobilisation. After one week of incubation, TA was decomposed and net C, N and P mineralization and net nitrification resembled that of the control (non-amended litter). CTs exhibited effects on net mineralization on longer terms, i.e. after several weeks of incubation until the end of the experiment (84 days). While net N and P mineralization were greatly reduced, net nitrification was only slightly affected. Most likely CTs formed complexes with organic N of the substrate thereby reducing net N mineralization, while such complexes were not involved in net nitrification processes. The reduction of net P mineralization is due to the lack of need for P by microbes when they cannot get access to N. The fact that decreasing amounts of extractable CTs were accompanied by increasing effects on mineralization processes with incubation time strongly suggests that CTs were incorporated into the litter in such a way that they were inextricable by the common solvents needed to measure tannins, such as for the Folin-Ciocalteu and HCl-butanol assays.

Climate change constitutes a serious threat for European heathlands as unlike other sources of damage, such as over-grazing, local remediation is not a possibility. Within the large pan-European projects, CLIMOOR and VULCAN, the effect of periodic drought and increased temperature were investigated in four heathland ecosystems along a geographical and climatic gradient across Europe. Fluorogenically labelled substrates for four enzymes (glucosidase, sulphatase, phosphatase, leucine amino peptidase) were used to measure extra-cellular enzyme activity in soil samples from each of the CLIMOOR sites. Microbial extra-cellular enzyme production is linked to microbial activity as well as soil physico-chemical properties, making soil enzymes one of the more reactive components of terrestrial ecosystems and potentially excellent indicators of soil microbial functional status and diversity. Across all sites and over all the substrates, organic matter content was exponentially, inversely related to enzyme activity. Although the increase in temperature produced by the CLIMOOR roofs was small (on average 0.9 °C), this was sufficient to increase enzyme activity in all sites (on average by 45%). The increase was within the range of seasonal variability at each of the sites. The effect of drought on enzyme activity was more pronounced in the Northern European sites than the southern European, and most moisture limited, site. This suggests that the effect of temperature increases may be observed across all regions; however, the soils of northern Europe may be more sensitive to changes in rainfall patterns than more moisture limited Southern European soils.

Climate change may affect the dynamics of ecosystems and the goods and services they provide. To investigate the consequences of warming and drought for the goods and services provided by different shrublands in various western European countries, an assessment was carried out using results of field manipulation experiments of the CLIMOOR and VULCAN projects. Goods and services of these shrublands mainly encompass biodiversity, various forms of recreation, conservation of culturally and historically important landscapes, groundwater as a drinking water source, and carbon sequestration. Warming of dry lowland heathlands in The Netherlands and Denmark increases nutrient availability, which may lead to grass encroachment reducing biodiversity and decreasing recreational values. Drought may reduce the chances of grass encroachment but increase the chances of disturbances to heather vegetation. Similarly, warming increases and drought decreases the chances of nitrate pollution to the groundwater, which is often used as a drinking water source. Warming of the upland heathland in the UK increases its productivity, which might enable higher grazing densities leading to improved agricultural production. However, complex interactions between heather and invading species may be affected. Furthermore, nitrate production is increased, which may lead to groundwater pollution. Under drought conditions, productivity decreases and agricultural production capacity drops. In the Mediterranean shrubland in Spain, both warming and drought led to a shift in the species composition of seedlings and recruitment, which might lead to a change in the plant community and a reduction in biodiversity. In the drought treatment, a decreasing soil carbon content may lead to a loss of biodiversity, recreational possibilities, and an increased threat of wildfires and erosion.

The value of nitrogen (N) field measurements for the calibration of parameters of the INCA nitrogen in catchment model is explored and quantified. A virtual catchment was designed by running INCA with a known set of parameters, and field "measurements" were selected from the model run output. Then, using these measurements and the Shuffled Complex Evolution Metropolis algorithm (SCEM-UA), four of the INCA model parameters describing N transformations in the soil were optimised, while the measurement uncertainty was increased in subsequent steps. Considering measurement uncertainty typical for N field studies, none of the synthesised datasets contained sufficient information to identify the model parameters with a reasonable degree of confidence. Parameter equifinality occurred, leading to considerable uncertainty in model parameter values and in modelled N concentrations and fluxes. Fortunately, combining the datasets in a multi-objective calibration was found to be effective in dealing with these equifinality problems. With the right choice of calibration measurements, multi-objective calibrations resulted in lower parameter uncertainty. The methodology applied in this study, using a virtual catchment free of model errors, is proposed as a useful tool foregoing the application of a N model or the design of a N monitoring program. For an already gauged catchment, a virtual study can provide a point of reference for the minimum uncertainty associated with a model application. When setting up a monitoring program, it can help to decide what and when to measure. Numerical experiments indicate that for a forested, N-saturated catchment, a fortnightly sampling of NO3 and NH4 concentrations in stream water may be the most cost-effective monitoring strategy. Keywords: INCA, nitrogen model, parameter uncertainty, multi-objective calibration, virtual catchment, experimental design

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.

Climate change may affect ecosystem functioning through increased temperatures or changes in precipitation patterns. Temperature and water availability are important drivers for ecosystem processes such as photosynthesis, carbon translocation, and organic matter decomposition. These climate changes may affect the supply of carbon and energy to the soil microbial population and subsequently alter decomposition and mineralization, important ecosystem processes in carbon and nutrient cycling. In this study, carried out within the cross-European research project CLIMOOR, the effect of climate change, resulting from imposed manipulations, on carbon dynamics in shrubland ecosystems was examined. We performed a 14C-labeling experiment to probe changes in net carbon uptake and allocation to the roots and soil compartments as affected by a higher temperature during the year and a drought period in the growing season. Differences in climate, soil, and plant characteristics resulted in a gradient in the severity of the drought effects on net carbon uptake by plants with the impact being most severe in Spain, followed by Denmark, with the UK showing few negative effects at significance levels of p ≤ 0.10. Drought clearly reduced carbon flow from the roots to the soil compartments. The fraction of the 14C fixed by the plants and allocated into the soluble carbon fraction in the soil and to soil microbial biomass in Denmark and the UK decreased by more than 60%. The effects of warming were not significant, but, as with the drought treatment, a negative effect on carbon allocation to soil microbial biomass was found. The changes in carbon allocation to soil microbial biomass at the northern sites in this study indicate that soil microbial biomass is a sensitive, early indicator of drought- or temperature-initiated changes in these shrubland ecosystems. The reduced supply of substrate to the soil and the response of the soil microbial biomass may help to explain the observed acclimation of CO2 exchange in other ecosystems.