Dr. Patrick Schleppi

Nitrogen (N) deposition has decreased in the last decades in Europe but in many cases remains higher than the critical loads, i.e., higher than what could be considered safe for biodiversity and ecosystem functioning. The main concerns about N deposition are eutrophication and acidification. In a long-term experiment (1994 to present) in a montane (1200 m a.s.l.) coniferous forest in Alptal, central Switzerland, we simulated increased N deposition by adding NH₄NO₃ to rainwater. This treatment consisted of an additional N input of 22 kg ha⁻¹ yr⁻¹ to the 12 kg ha⁻¹ yr⁻¹ ambient bulk deposition rate or 17 kg N ha⁻¹ yr⁻¹ throughfall rate. The treatment was applied simultaneously to a small catchment area and to plots in a replicated block experiment (n=5). The site has a carbonate-rich parent material and is thus not particularly at risk of acidification. Nevertheless, we examined soil acidification as affected both by ambient and experimentally increased N deposition. In the 2.5 decades since the beginning of the study, nitrate (NO₃^-) and especially sulfate (SO₄^2-) concentrations decreased in precipitation, while pH increased by slightly more than 1 unit. In the same time period a reduction in pH of the soil was measured. The exchangeable acidity in the soil increased, especially in the N-addition treatment. This was mainly observed on small mounds because the drier mounds are less well buffered than wet depressions. This trend, however, was limited in time, as exchangeable acidity later declined again to reach values not much higher than 26 years before. This was also the case in the N-addition treatment and can be considered a progressive recovery mainly due to the reduced acid inputs and, at this site with a carbonate-rich subsoil, to the biological cycling of base cations. The pH of the runoff from the experimental catchments decreased by 0.3 units, both in the control and under N addition. Decreasing Ca²⁺ and increasing Al³⁺ and Fe²⁺ concentrations in runoff also show that the recovery observed in the exchangeable soil acidity is not yet able to stop the slow acidification of water leaving the catchments. However, with the runoff water pH remaining above 7, this trend is not alarming for water quality or for the health of water bodies. Future monitoring will be necessary to see if and when a recovery takes place in the soil and runoff pH.

Soil matric potential quantifies water availability in soils. Low soil matric potentials are difficult to measure with most in situ techniques. This is also the case for the widely-used dielectric MPS-2 sensor. This probe determines matric potential indirectly from the measured water content in its porous sensor ceramics using dielectric permittivity as a proxy for water content. The accuracy of MPS-2 readings was analyzed in desiccation experiments using 13 soils with different texture and organic carbon content and a WP4C dewpoint potentiometer as reference instrument. Further, it was explored whether observed inaccuracies relate to sensor calibration, confounding dielectric effects, or the water release characteristics of the sensor ceramics. Above -1000 kPa, the MPS-2 readings were accurate in all tested soils with a mean deviation of 3% to the reference values. Below -1000 kPa, MPS-2 readings were increasingly higher than the reference in all tested soils, but the deviation from the reference depended on soil type. Poor factory calibration of the sensors, soil texture dependent differences of water flow at the soil-ceramic interface, and dielectric effects are supposed to be the main reasons for the low and soil-type-specific MPS-2 sensor accuracy in dry soils. Nevertheless, the high consistency of the MPS-2 readings allowed us to derive soil-type-specific equations to improve the accuracy of measurements to values as low as -4000 kPa. We recommend applying the equations to any MPS-2 readings below ~ -1000 kPa to obtain more reliable data and thus an improved insight into the role of soil water in ecosystems.

Climate change exposes ecosystems to strong and rapid changes in their environmental boundary conditions mainly due to the altered temperature and precipitation patterns. It is still poorly understood how fast interlinked ecosystem processes respond to altered environmental conditions, if these responses occur gradually or suddenly when thresholds are exceeded, and if the patterns of the responses will reach a stable state. We conducted an irrigation experiment in the Pfynwald, Switzerland from 2003–2018. A naturally dry Scots pine (Pinus sylvestris L.) forest was irrigated with amounts that doubled natural precipitation, thus releasing the forest stand from water limitation. The aim of this study was to provide a quantitative understanding on how different traits and functions of individual trees and the whole ecosystem responded to increased water availability, and how the patterns and magnitudes of these responses developed over time. We found that the response magnitude, the temporal trajectory of responses, and the length of initial lag period prior to significant response largely varied across traits. We detected rapid and stronger responses from aboveground tree traits (e.g., tree-ring width, needle length, and crown transparency) compared to belowground tree traits (e.g., fine-root biomass). The altered aboveground traits during the initial years of irrigation increased the water demand and trees adjusted by increasing root biomass during the later years of irrigation, resulting in an increased survival rate of Scots pine trees in irrigated plots. The irrigation also stimulated ecosystem-level foliar decomposition rate, fungal fruit body biomass, and regeneration abundances of broadleaved tree species. However, irrigation did not promote the regeneration of Scots pine trees, which are reported to be vulnerable to extreme droughts. Our results provide extensive evidence that tree- and ecosystem-level responses were pervasive across a number of traits on long-term temporal scales. However, after reaching a peak, the magnitude of these responses either decreased or reached a new stable state, providing important insights into how resource alterations could change the system functioning and its boundary conditions.

Human demand for food and energy has led to significant changes in the level of reactive nitrogen (N) released to the atmosphere and then deposited in the biosphere. This study aimed to investigate the impact of the deposition of artificial N on the forest floor and on the soil chemical properties in an oak (Quercus castaneifolia C.A. Mey.) plantation in northern Iran. Twelve plots of 200 m² (20 m × 10 m) were set up in the study area. Four N treatments were considered: zero (control), 50 (low), 100 (medium), and 150 (high) kg N·ha^-1·year^-1. N in the form of NH₄NO₃ solution was manually sprayed onto the understory plots monthly for 1 year. The total N, phosphorus (P), potassium (K), and organic carbon (OC) of the forest floor were measured. Soil N, available P, available K, pH, EC (electrical conductivity), OC, microbial biomass C (MBC), and urease enzyme activity were measured in the 0-10 cm depth. The concentration of total N and P of the forest floor was significantly higher in the high-N treatment. The total concentration of N (+36%), the urease activity (+44%), and EC (+12%) of soil increased with raising the high-N treatment compared to the control, but the MBC (-20%), available P (-28%), and available K (-15%) were significantly reduced in the high-N treatment. Our results were obtained with simulated deposition rates that exceed ambient fluxes, but ambient N deposition is nevertheless high in our study area.

Wie viel Licht ist im Wald? Wie gross ist die Blattfläche des Waldes? Um diese Fragen in einem Bestand quantitativ beantworten zu können, bietet die hemispärische Fotografie eine bewährte Möglichkeit.

Previous studies have evaluated how changes in atmospheric nitrogen (N) inputs and climate affect stream N concentrations and fluxes, but none have synthesized data from sites around the globe. We identified variables controlling stream inorganic N concentrations and fluxes, and how they have changed, by synthesizing 20 time series ranging from 5 to 51 years of data collected from forest and grassland dominated watersheds across Europe, North America, and East Asia and across four climate types (tropical, temperate, Mediterranean, and boreal) using the International Long-Term Ecological Research Network. We hypothesized that sites with greater atmospheric N deposition have greater stream N export rates, but that climate has taken a stronger role as atmospheric deposition declines in many regions of the globe. We found declining trends in bulk ammonium and nitrate deposition, especially in the longest time-series, with ammonium contributing relatively more to atmospheric N deposition over time. Among sites, there were statistically significant positive relationships between (1) annual rates of precipitation and stream ammonium and nitrate fluxes and (2) annual rates of atmospheric N inputs and stream nitrate concentrations and fluxes. There were no significant relationships between air temperature and stream N export. Our long-term data shows that although N deposition is declining over time, atmospheric N inputs and precipitation remain important predictors for inorganic N exported from forested and grassland watersheds. Overall, we also demonstrate that long-term monitoring provides understanding of ecosystems and biogeochemical cycling that would not be possible with short-term studies alone.

Leaf morphological traits (LMTs) of forest trees have been observed to vary across space and species. However, long-term records of LMTs are scarce, due to a lack of measurements and systematic leaf archives. This leaves a large gap in our understanding of the temporal dynamics and drivers of LMT variations, which may help us understand tree acclimation strategies. In our study, we used long-term LMT measurements from foliar material collections of European beech (Fagus sylvatica) and Norway spruce (Picea abies), performed every second year from 1995 to 2019 on the same trees within the Swiss Long-term Forest Ecosystem Research Program LWF. The 11 study plots (6 beech, 4 spruce, and 1 mixed) are distributed along gradients of elevation (485-1,650 m a.s.l.), mean annual precipitation (935-2142 mm), and mean annual temperature (3.2-9.8°C). The investigated LMTs were (i) leaf or needle mass, (ii) leaf area or needle length, and (iii) leaf mass per area or needle mass per length. We combined this unique data set with plot variables and long-term data on potential temporal drivers of LMT variations, including meteorological and tree trait data. We used univariate linear regressions and linear mixed-effects models to identify the main spatial and temporal drivers of LMT variations, respectively. For beech LMTs, our temporal analysis revealed effects of mast year and crown defoliation, and legacy effects of vapor pressure deficit and temperature in summer and autumn of the preceding year, but no clear long-term trend was observed. In contrast, spruce LMTs were mainly driven by current-year spring conditions, and only needle mass per length showed a decreasing long-term trend over the study period. In temporal models, we observed that LMTs of both species were influenced by elevation and foliar nutrient concentrations, and this finding was partly confirmed by our spatial analyses. Our results demonstrate the importance of temporal analysis for determining less recognized drivers and legacy effects that influence LMTs, which are difficult to determine across space and species. The observed differences in the temporal drivers of beech and spruce LMTs suggest differences in the adaptation and acclimation potential of the two species.

Oxidative weathering of sedimentary rocks plays an important role in the global carbon cycle. Rhenium (Re) has been proposed as a tracer of rock organic carbon (OC_petro) oxidation. However, the sources of Re and its mobilization by hydrological processes remain poorly constrained. Here, we examine dissolved Re as a function of water discharge, using samples collected from three alpine catchments that drain sedimentary rocks in Switzerland (Erlenbach and Vogelbach) and Colorado, USA (East River). The Swiss catchments reveal a higher dissolved Re flux in the catchment with higher erosion rates, but have similar [Re]/[Na⁺] and [Re]/[SO₄^2-] ratios, which indicate a dominance of Re from OC_petro. Despite differences in rock type and hydro-climatic setting, the three catchments have a positive correlation between river water [Re]/[Na⁺] and [Re]/[SO₄^2-] and water discharge. We propose that this reflects preferential routing of Re from a near-surface, oxidative weathering zone. The observations support the use of Re as a proxy to trace rock-organic carbon oxidation, and suggest it may be a hydrological tracer of vadose zone processes. We apply the Re proxy and estimate CO₂ release by OC_petro oxidation of 5.7 ^+6.6/_-2.0 tC km^-2 yr^-1 for the Erlenbach. The overall weathering intensity was ∼40%, meaning that the corresponding export of unweathered OC_petro in river sediments is large, and the findings call for more measurements of OC_petro oxidation in mountains and rivers as they cross floodplains.

The stable isotope ¹⁵N is an extremely useful tool for studying the nitrogen (N) cycle of terrestrial ecosystems. The affordability of isotope-ratio mass spectrometry has increased in the last decades and routine measurements of δ¹⁵N with an accuracy better than 1‰ are now easily achieved. Except perhaps for wood, which has a very high C/N ratio, isotope analysis of samples is, thus, no longer the main challenge in measuring the partitioning of ¹⁵N used as tracer in ecosystem studies. The central aim of such experiments is to quantitatively determine the fate of N after it enters an ecosystem, mainly as fertilizer, as atmospheric deposition or as plant litter. By measuring how much of this incoming N goes into different ecosystem pools, inferences can be made about the entire N cycle. Sample collection and preparation can be tedious work. Optimizing sampling schemes is thus an important aspect in the application of ¹⁵N in ecosystem research and can be helpful for obtaining a high precision of the results with the available manpower and budget. In this contribution, we combine statistical and practical considerations and give recommendations for the design of labeling experiments and also for assessments of natural ¹⁵N abundance. In particular, we discuss soil, vegetation and water sampling. We additionally address the most common questions arising during the calculation of tracer partitioning, and we provide some examples of the interpretation of experimental results.

Light availability below and within forest canopies governs many biological processes. Its estimation, however, is often time consuming, especially in complex canopy structures where vertical profiles of light transmission need to be known to estimate plant performance, habitat suitability or biophysical properties across the canopy. To overcome the technical limitations of assessing vertical canopy space by ground-based systems like ropes, towers or cranes, we developed a prototype to take hemispherical photographs (HP) from an unmanned aerial vehicle (UAV). With the new airborne system, we aimed to assess the differences of vertical light availability profiles from bottom to the top of the canopy in even-aged, uneven-aged and coppice-with-standards forests. Measured vertical light profiles typically followed a sigmoid shape that differed markedly depending on canopy gap size and forest structure. The results showed important variations in the transmission of sunlight at different heights within forest canopies, highlighting the importance of vertical canopy stratification when considering radiation transfer, plant area index (PAI), leaf area index (LAI) and light availability. To further cover cases where branches hinder UAV access to dense canopy spaces, we present a workflow to estimate light availability from photogrammetrically reconstructed high-resolution vegetation height models (VHM) and validate the outputs against HP. We conclude that VHMs based on high-resolution aerial photography, photogrammetry and a structure-from-motion algorithm (SfM) deliver a solid foundation to derive a wide range of canopy metrics. With an appropriate VHM point density, direct or beam light index (BLI), diffuse light index (DLI), gap or global light index (GLI) and PAI in mixed broadleaved forests can be reliably calculated. The novel development of UAV-based HP and the workflow to calculate light regimes from VHMs thus open a wide range of new applications in forest and agricultural research and management.

Soil appears to play a key role in the response of the forest ecosystems to N deposition. Twenty years of experimental moderate N addition in a sub-alpine forest increased nitrate leaching, but the soil immobilized most of the N input, gradually decreasing the C:N ratio. Exchangeable and microbial N were only slightly affected, but denitrification and N₂O production were increased and soil respiration tended to be reduced while soil microbial communities were remarkably resistant. It is assumed that these changes at the process level are related to the soil microbiome, but soil microbial communities have not been assessed so far at lower taxonomical resolution in this long-term experiment. The aim of this study is to understand the underlying causes of the results obtained so far by assessing how N treatment affects the soil microbiome at different soil depths. We analyzed bacterial and fungal diversity and community structures using Illumina MiSeq sequencing and quantified the responses of the N cycling communities to elevated N loads by quantitative PCR. The microbial functions were assessed by respiration, N mineralization, and potential nitrification. Bacterial and fungal α-diversity, observed richness and Shannon diversity index, remained unchanged upon N addition. Multivariate statistics showed shifts in the structures of fungal but not bacterial communities with N load, while the changes were minor. Differences in the community compositions associated with the N treatment were mainly observed at a lower taxonomical level. We found several fungal OTUs in particular genera such as the ectomycorrhizal fungi Hydnum, Piloderma, Amanita, and Tricholoma that decreased significantly with increased N-loads. We conclude that long-term moderate N addition at this forest site did not strongly affect the soil microbiome (which remained remarkably resistant) and its functioning.

Climate change and the expected increase in frequency of dry summers are likely to affect the growth of important tree species. We investigated relationships between morphological and chemical leaf traits of European beech (Fagus sylvatica L.) and environmental factors along a water availability gradient consisting of 12 sites located throughout Switzerland. We found that leaf dry mass and leaf area tended to decrease with increasing long-term mean annual precipitation (MAP) and actual to potential transpiration (AT/PT), two correlated variables that form the water availability gradient. These results contrast with those of several other studies, and might be explained by favorable temperature and humidity conditions during leaf formation in spring at the dry sites. Although the relationship was not linear over the whole gradient, the drier sites were characterized by beech with a lower specific leaf area, along with higher foliar potassium (K) and lower foliar nitrogen (N) concentrations. These patterns likely reflect strategies developed as an adaptation to reduced water availability, but they also result from variation in the availability of nutrients in soil across our sites. In the case of N, there are indications that atmospheric deposition plays a role in foliar concentrations.

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.

To accurately capture the impacts of nitrogen (N) on the land carbon (C) sink in Earth system models, model responses to both N limitation and ecosystem N additions (e.g., from atmospheric N deposition and fertilizer) need to be evaluated. The response of the land C sink to N additions depends on the fate of these additions: that is, how much of the added N is lost from the ecosystem through N loss pathways or recovered and used to increase C storage in plants and soils. Here, we evaluate the C–N dynamics of the latest version of a global land model, the Community Land Model version 5 (CLM5), and how they vary when ecosystems have large N inputs and losses (i.e., an open N cycle) or small N inputs and losses (i.e., a closed N cycle). This comparison allows us to identify potential improvements to CLM5 that would apply to simulated N cycles along the open-to-closed spectrum. We also compare the short- (< 3 years) and longer-term (5–17 years) N fates in CLM5 against observations from 13 long-term ¹⁵N tracer addition experiments at eight temperate forest sites. Simulations using both open and closed N cycles overestimated plant N recovery following N additions. In particular, the model configuration with a closed N cycle simulated that plants acquired more than twice the amount of added N recovered in ¹⁵N tracer studies on short timescales (CLM5: 46±12 %; observations: 18±12 %; mean across sites ±1 standard deviation) and almost twice as much on longer timescales (CLM5: 23±6 %; observations: 13±5 %). Soil N recoveries in simulations with closed N cycles were closer to observations in the short term (CLM5: 40±10 %; observations: 54±22 %) but smaller than observations in the long term (CLM5: 59±15 %; observations: 69±18 %). Simulations with open N cycles estimated similar patterns in plant and soil N recovery, except that soil N recovery was also smaller than observations in the short term. In both open and closed sets of simulations, soil N recoveries in CLM5 occurred from the cycling of N through plants rather than through direct immobilization in the soil, as is often indicated by tracer studies. Although CLM5 greatly overestimated plant N recovery, the simulated increase in C stocks to recovered N was not much larger than estimated by observations, largely because the model's assumed C:N ratio for wood was nearly half that suggested by measurements at the field sites. Overall, results suggest that simulating accurate ecosystem responses to changes in N additions requires increasing soil competition for N relative to plants and examining model assumptions of C:N stoichiometry, which should also improve model estimates of other terrestrial C–N processes and interactions.

Forest soils harbor diverse microbial communities responsible for the cycling of elements including carbon (C), nitrogen (N), and phosphorus (P). Conversely, anthropogenic N deposition can negatively feed back on soil microbes and reduce soil organic matter (SOM) decomposition. Mechanistically, this can include reductions of decomposer biomass, especially fungi, and decreases in activities of lignin-modifying enzyme (LMEs). Moreover, N inputs can lower resource C:N and thus decrease the C:N imbalance between microbial decomposers and their resources. As a result, microbially-mediated decomposition might slow down, resulting in larger SOM pools with consequences for ecosystem nutrition and climate regulation. Here, we studied the long-term impact of experimental N addition on soil microbes and microbially-mediated decomposition in two coniferous forests in Switzerland and Denmark. We measured microbial biomass C and N, phospholipid fatty acid (PLFA) biomarkers and potential enzyme activities related to C, N, and P acquisition along the topsoil profile (0-30 cm). In particular, we investigated shifts in microbial C:N homeostasis and relative C:N:P limitation. Contrary to prevailing theory, microbial biomass and community composition were remarkably resistant against two decades of 750 and 1,280 kg ha^-1 of cumulative N inputs at the Swiss and Danish site, respectively. While N reduced fungal-specific PLFAs and lowered fungi-to-bacteria (F:B) ratios in some (mainly organic) horizons where soil organic carbon (SOC) has accumulated, it increased F:B ratios in other (mainly mineral) horizons where SOC has declined. We did not find a consistent reduction of LME activities in response to N. Rather, relationships between LME activities and SOC concentrations were largely unaffected by N addition. This questions prevalent theories of lignin decomposition and SOC storage under elevated N inputs. By using C:N stoichiometry, we further show that microbial communities responded in part nonhomeostatically to decreasing resource C:N, in addition to a likely increase in their carbon use efficiency and a decrease in nitrogen use efficiency. While the expected increased allocation to C- and decreased allocation to N-acquiring enzymes was not found, microbial investment in P acquisition (acid phosphatase activity) increased in the nutrient-poor Podzol (but not in the nutrient-rich Gleysol). Enzyme vector analysis showed decreasing C but increasing P limitation of soil microbial communities at both sites. We conclude that simulated N deposition led to physiological adaptations of soil microbial communities across the topsoil profile in two independent experiments, with long-term implications for tree nutrition and SOC sequestration. However, we expect that microbial adaptations are not endless and may reach a tipping point when ecosystems experience nitrogen saturation.

Nitrogen (N) inputs from atmospheric deposition can increase soil organic carbon (SOC) storage in temperate and boreal forests, thereby mitigating the adverse effects of anthropogenic CO₂; emissions on global climate. However, direct evidence of N-induced SOC sequestration from low-dose, long-term N addition experiments (that is, addition of < 50 kg N ha^–1 y^–1; for > 10 years) is scarce worldwide and virtually absent for European temperate forests. Here, we examine how tree growth, fine roots, physicochemical soil properties as well as pools of SOC and soil total N responded to 20 years of regular, low-dose N addition in two European coniferous forests in Switzerland and Denmark. At the Swiss site, the addition of 22 kg N ha^–1 y^–1 (or 1.3 times throughfall deposition) stimulated tree growth, but decreased soil pH and exchangeable calcium. At the Danish site, the addition of 35 kg N ha^–1 y^–1 (1.5 times throughfall deposition) impaired tree growth, increased fine root biomass and led to an accumulation of N in several belowground pools. At both sites, elevated N inputs increased SOC pools in the moderately decomposed organic horizons, but decreased them in the mineral topsoil. Hence, long-term N addition led to a vertical redistribution of SOC pools, whereas overall SOC storage within 30 cm depth was unaffected. Our results imply that an N-induced shift of SOC from older, mineral-associated pools to younger, unprotected pools might foster the vulnerability of SOC in temperate coniferous forest soils.

The response of trees to increasing atmospheric CO₂ concentrations is often mediated by the availability of nutrients. However, little is known about the influence of CO₂ enrichment on nutrient availability in forests with mature trees. We studied processes in the soil under five 35-m-tall Norway spruce trees (Picea abies) in NW Switzerland that were exposed to a mean CO₂ concentration of 550 ppm for 5 growing seasons using free air CO₂ enrichment (FACE). We compared them with values from the soil under five control trees. Ceramic suction cups were installed in the soil under each tree to collect soil solution, and ion-exchange resin bags were buried in the soil to absorb ammonium and nitrate. Soil cores taken at the end of the experiment were used to measure the gross production of ammonium and nitrate by the ¹⁵N dilution technique. Although temporally and spatially variable, the nitrate concentration was higher in the soil solution under CO₂-enriched trees. This effect was reflected in the resin bag extracts, which additionally indicated a trend of increased ammonium availability. Dissolved reduced N concentration (mainly dissolved organic N), however, was lower in the soil solution under CO₂-enriched trees. K and Mg, and to a lesser extent Na and sulfate concentrations increased in the soil solution. P concentrations mostly remained below the detection limit. In spite of the higher concentrations of nitrate in soil extracts, gross N mineralization and nitrification rates were not affected by FACE. Needles from CO₂-enriched trees contained slightly more N. No difference was observed for other nutrients. Overall, these results support the hypothesis of a priming effect, i.e., that FACE led to the production of more root exudates, which in turn stimulated soil biological activity, including mineralization, over a time-span of at least several years. However, these tall trees showed no growth response to elevated CO₂; hence, they gained no advantage from increased nitrate in the soil solution, presumably owing to other growth constraints including P and Mg availability.

Atmospheric nitrogen (N) deposition in terrestrial ecosystems is difficult to quantify, especially in forests. In this study, we compared three approaches for determining the wet and dry deposition of nitrogen (total deposition) at 17 intensively monitored forest sites in Switzerland. Specifically, we considered approaches based on: 1) measurements of bulk deposition and throughfall in 2014 (throughfall method); 2) measurements of bulk deposition and measurements of air concentrations of ammonia (NH₃) and nitrogen dioxide (NO₂), to which deposition velocities were applied, also in 2014 (inferential method); and 3) a model developed for Switzerland at a high spatial resolution, run for the five-year period 2013–2017 (emission based model). In addition, changes over two decades were assessed using continuous measurements of throughfall and bulk deposition. Further, air concentrations of NH₃ and NO₂ measured in 2014 were compared with concentrations measured in 2000 at 10 of the sites. The three approaches generally yielded comparable estimates of total deposition, with some notable differences at some sites. For both the model and the inferential method, uncertainties were related to the deposition velocities that were applied to air concentrations of N compounds, especially for NH₃. The throughfall method provided a minimum estimate of the total N deposition in the forest, but the fraction of the deposited N that is directly taken up by the canopy remained difficult to quantify. Nitrogen deposition has decreased since the mid-1990s at the majority of the sites. However, deposition of the reduced forms of N seems to have stagnated at the local level. Furthermore, N deposition is still too high in comparison with the range of empirical critical loads of N (CLN). The minimum deposition estimated from throughfall exceeds the lower limit of CLN at all sites except those in the Central Alps. Deposition estimated with the model and the inferential method exceeds the lower limit of CLN at all sites and even exceeds the upper limit at several locations.

Light is a key driver of forest biodiversity and functioning. Light regimes beneath tree canopies are mainly driven by the solar angle, topography, and vegetation structure, whose three‐dimensional complexity creates heterogeneous light conditions that are challenging to quantify, especially across large areas. Remotely sensed canopy structure data from airborne laser scanning (ALS) provide outstanding opportunities for advancement in this respect. We used ALS point clouds and a digital terrain model to produce hemispherical photographs from which we derived indices of nondirectional diffuse skylight and direct sunlight reaching the understory. We validated our approach by comparing the performance of these indices, as well as canopy closure (CCl) and canopy cover (CCo), for explaining the light conditions experienced by forest plant communities, as indicated by the Landolt indicator values for light (L_light) from 43 vegetation surveys along an elevational gradient. We applied variation partitioning to analyze how the independent and joint statistical effects of light, macroclimate, and soil on the spatial variation in plant species composition (i.e., turnover, Simpson dissimilarity, β_SIM) depend on light approximation methodology. Diffuse light explained L_light best, followed by direct light, CCl and CCo (R² = .31, .23, .22, and .22, respectively). The combination of diffuse and direct light improved the model performance for β_SIMcompared with CCl and CCo (R² = .30, .27 and .24, respectively). The independent effect of macroclimate on β_SIM dropped from an R² of .15 to .10 when diffuse light and direct light were included. The ALS methods presented here outperform conventional approximations of below‐canopy light conditions, which can now efficiently be quantified along entire horizontal and vertical forest gradients, even in topographically complex environments such as mountains. The effect of macroclimate on forest plant communities is prone to be overestimated if local light regimes and associated microclimates are not accurately accounted for.