Soil warming opens the nitrogen cycle at the alpine treeline.

Climate warming may alter ecosystem nitrogen (N) cycling by accelerating N transformations in the soil, and changes may be especially pronounced in cold regions characterized by N-poor ecosystems. We investigated N dynamics across the plant-soil continuum during 6 years of experimental soil warming (2007-2012; +4 °C) at a Swiss high-elevation treeline site (Stillberg, Davos; 2180 m a.s.l.) featuring Larix decidua and Pinus uncinata. In the soil, we observed considerable increases in the NH4+ pool size in the first years of warming (by >50%), but this effect declined over time. In contrast, dissolved organic nitrogen (DON) concentrations in soil solutions from the organic layer increased under warming, especially in later years (maximum of +45% in 2012), suggesting enhanced DON leaching from the main rooting zone. Throughout the experimental period, foliar N concentrations showed species-specific but small warming effects, whereas δ15 N values showed a sustained increase in warmed plots that was consistent for all species analysed. The estimated total plant N pool size at the end of the study was greater (+17%) in warmed plots with Pinus but not in those containing Larix, with responses driven by trees. Irrespective of plot tree species identity, warming led to an enhanced N pool size of Vaccinium dwarf shrubs, no change in that of Empetrum hermaphroditum (dwarf shrub) and forbs, and a reduction in that of grasses, nonvascular plants, and fine roots. In combination, higher foliar δ15 N values and the transient response in soil inorganic N indicate a persistent increase in plant-available N and greater cumulative plant N uptake in warmer soils. Overall, greater N availability and increased DON concentrations suggest an opening of the N cycle with global warming, which might contribute to growth stimulation of some plant species while simultaneously leading to greater N losses from treeline ecosystems and possibly other cold biomes.

Reassessment of the NH4 NO3 thermal decomposition technique for calibration of the N2 O isotopic composition.

RATIONALE: In the last few years, the study of N2 O site-specific nitrogen isotope composition has been established as a powerful technique to disentangle N2 O emission pathways. This trend has been accelerated by significant analytical progress in the field of isotope ratio mass spectrometry (IRMS) and more recently quantum cascade laser absorption spectroscopy (QCLAS). METHODS: The ammonium nitrate (NH4 NO3 ) decomposition technique provides a strategy to scale the 15 N site-specific (SP ≡ Î´15 Nα - δ15 Nß ) and bulk (δ15 Nbulk  = (δ15 Nα  + Î´15 Nß )/2) isotopic composition of N2 O against the international standard for the 15 N/14 N isotope ratio (AIR-N2 ). Within the current project 15 N fractionation effects during thermal decomposition of NH4 NO3 on the N2 O site preference were studied using static and dynamic decomposition techniques. RESULTS: The validity of the NH4 NO3 decomposition technique to link NH4+ and NO3- moiety-specific δ15 N analysis by IRMS to the site-specific nitrogen isotopic composition of N2 O was confirmed. However, the accuracy of this approach for the calibration of δ15 Nα and δ15 Nß values was found to be limited by non-quantitative NH4 NO3 decomposition in combination with substantially different isotope enrichment factors for the conversion of the NO3- or NH4+ nitrogen atom into the α or ß position of the N2 O molecule. CONCLUSIONS: The study reveals that the completeness and reproducibility of the NH4 NO3 decomposition reaction currently confine the anchoring of N2 O site-specific isotopic composition to the international isotope ratio scale AIR-N2 . The authors suggest establishing a set of N2 O isotope reference materials with appropriate site-specific isotopic composition, as community standards, to improve inter-laboratory compatibility. Copyright © 2016 John Wiley & Sons, Ltd.

Effect of irrigation on needle morphology, shoot and stem growth in a drought-exposed Pinus sylvestris forest.

In Valais, Switzerland, Scots pines (Pinus sylvestris L.) are declining, mainly following drought. To assess the impact of drought on tree growth and survival, an irrigation experiment was initiated in 2003 in a mature pine forest, approximately doubling the annual precipitation. Tree crown transparency (lack of foliage) and leaf area index (LAI) were annually assessed. Seven irrigated and six control trees were felled in 2006, and needles, stem discs and branches were taken for growth analysis. Irrigation in 2004 and 2005, both with below-average precipitation, increased needle size, area and mass, stem growth and, with a 1-year delay, shoot length. This led to a relative decrease in tree crown transparency (-14%) and to an increase in stand LAI (+20%). Irrigation increased needle length by 70%, shoot length by 100% and ring width by 120%, regardless of crown transparency. Crown transparency correlated positively with mean needle size, shoot length and ring width and negatively with specific leaf area. Trees with high crown transparency (low growth, short needles) experienced similar increases in needle mass and growth with irrigation than trees with low transparency (high growth, long needles), indicating that seemingly declining trees were able to 'recover' when water supply became sufficient. A simple drought index before and during the irrigation explained most of the variation found in the parameters for both irrigated and control trees.

Estimating below-canopy light regimes using airborne laser scanning: An application to plant community analysis.

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, ß 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 ß SIM compared with CCl and CCo (R2  = .30, .27 and .24, respectively). The independent effect of macroclimate on ß SIM dropped from an R 2 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.

Dynamics of atmospheric nitrogen deposition in a temperate calcareous forest soil.

In temperate forest ecosystems, soil acts as a major sink for atmospheric N deposition. A (15)N labeling experiment in a hardwood forest on calcareous fluvisol was performed to study the processes involved. Low amounts of ammonium ((15)NH(4)(+)) or nitrate ((15)NO(3)(-)) were added to small plots. Soil samples were taken after periods ranging from 1 h to 1 yr. After 1 d, the litter layer retained approximately 28% of the (15)NH(4)(+) tracer and 19% of (15)NO(3)(-). The major fraction of deposited N went through the litter layer to reach the soil within the first hours following the tracer application. During the first day, a decrease in extractable (15)N in the soil was observed ((15)NH(4)(+): 50 to 5%; (15)NO(3)(-): 60 to 12%). During the same time, the amount of microbial (15)N remained almost constant and the (15)N immobilized in the soil (i.e., total (15)N recovered in the bulk soil minus extractable (15)N minus microbial (15)N) also decreased. Such results can therefore be understood as a net loss of (15)N from the soil. Such N loss is probably explained by NO(3)(-) leaching, which is enhanced by the well-developed soil structure. We presume that the N immobilization mainly occurs as an incorporation of deposited N into the soil organic matter. One year after the (15)N addition, recovery rates were similar and approximately three-quarters of the deposited N was recovered in the soil. We conclude that the processes relevant for the fate of atmospherically deposited N take place rapidly and that N recycling within the microbes-plants-soil organic matter (SOM) system prevents further losses in the long term.

Responses of soil Collembola to long-term atmospheric CO2 enrichment in a mature temperate forest.

Responses of Collembola to 7 years of CO(2) enrichment (550 ppm) in a Swiss free-air CO(2) enrichment (FACE) experiment in a forest with 80- to 120-year-old trees were investigated in this study. Contrary to our expectations, increased CO(2) caused a significant decrease in Collembola numbers, including a significant decrease in euedaphic Collembola. Increased CO(2), however, did not affect community group richness. Collembola biomass was not significantly changed by CO(2) enrichment, regardless of whether it was considered in terms of the total community, life-strategy groups, or individual species (with an exception of Mesaphorura krausbaueri). The reason for this is that CO(2) enrichment caused a general increase in individual body size, which compensated for reduced abundances. The results are consistent with the idea that the rhizosphere is important for soil fauna, and the combination of reduced fine root growth and increased soil moisture might trigger a reduction in Collembola abundance.

Growth enhancement of Picea abies trees under long-term, low-dose N addition is due to morphological more than to physiological changes.

Human activities have drastically increased nitrogen (N) inputs into natural and near-natural terrestrial ecosystems such that critical loads are now being exceeded in many regions of the world. This implies that these ecosystems are shifting from natural N limitation to eutrophication or even N saturation. This process is expected to modify the growth of forests and thus, along with management, to affect their carbon (C) sequestration. However, knowledge of the physiological mechanisms underlying tree response to N inputs, especially in the long term, is still lacking. In this study, we used tree-ring patterns and a dual stable isotope approach (δ(13)C and δ(18)O) to investigate tree growth responses and the underlying physiological reactions in a long-term, low-dose N addition experiment (+23 kg N ha(-1) a(-1)). This experiment has been conducted for 14 years in a mountain Picea abies (L.) Karst. forest in Alptal, Switzerland, using a paired-catchment design. Tree stem C sequestration increased by ∼22%, with an N use efficiency (NUE) of ca. 8 kg additional C in tree stems per kg of N added. Neither earlywood nor latewood δ(13)C values changed significantly compared with the control, indicating that the intrinsic water use efficiency (WUE(i)) (A/g(s)) did not change due to N addition. Further, the isotopic signal of δ(18)O in early- and latewood showed no significant response to the treatment, indicating that neither stomatal conductance nor leaf-level photosynthesis changed significantly. Foliar analyses showed that needle N concentration significantly increased in the fourth to seventh treatment year, accompanied by increased dry mass and area per needle, and by increased tree height growth. Later, N concentration and height growth returned to nearly background values, while dry mass and area per needle remained high. Our results support the hypothesis that enhanced stem growth caused by N addition is mainly due to an increased leaf area index (LAI). Higher LAI implies that more photosynthetically active radiation is absorbed and therefore canopy-level photosynthesis is increased. We conclude that models assuming that N deposition increases tree growth through higher leaf-level photosynthesis may be mechanistically inaccurate, at least in forest canopies that are not (yet) completely closed.

Negative responses of Collembola in a forest soil (Alptal, Switzerland) under experimentally increased N deposition.

The response of specific groups of organisms, like Collembola to atmospheric nitrogen (N) deposition is still scarcely known. We investigated the Collembola community in a subalpine forest (Alptal, Switzerland) as subjected for 12 years to an experimentally increased N deposition (+25 on top of ambient 12 kg N ha(-1) year(-1)). In the 0-5 cm soil layer, there was a tendency of total Collembola densities to be lower in N-treated than in control plots. The density of Isotomiella minor, the most abundant species, was significantly reduced by the N addition. A tendency of lower Collembola group richness was observed in N-treated plots. The Density-Group index (dDG) showed a significant reduction of community diversity, but the Shannon-Wiener index (H') was not significantly affected by the N addition. The Collembola community can be considered as a bioindicator of N inputs exceeding the biological needs, namely, soil N saturation.

Citric acid traps to replace sulphuric acid in the ammonia diffusion of dilute water samples for 15N analysis.

The analysis of 15N in aqueous samples requires the concentration of dissolved nitrogen (N) into a small volume that can be analysed by mass spectrometry. This is conveniently achieved by the NH3 diffusion technique, where NH4+ is captured on small acidified filters enclosed in PTFE. NO3- can be analysed the same way by reducing it to NH4+ with Devarda's alloy. H2SO4 is commonly used for the acidification of the filters. During combustion, however, this acid leads to the production of SO2 and elemental sulphur, which both have detrimental effects on the mass spectrometer. We propose here to replace H2SO4 with citric acid because it is combusted completely to CO2 and H2O in the elemental analyser before entering the mass spectrometer. Citric acid was found to give comparable results in terms of N recovery and 15N values, both for NH4+ and for NO3- samples. Blank samples revealed that N contamination was slightly lower using citric instead of sulphuric acid as acidifier of the glass filters. NH4+ samples first concentrated over cation-exchange columns were strongly acidic and several methods were tested to raise the pH for the subsequent diffusion. These samples gave incomplete N recoveries, but this problem was independent of the acid used on the filters and of the final pH of the sample. Complete recovery was achieved only by increasing the volume of the eluate from the columns. Citric acid can thus generally be recommended instead of H2SO4 for ammonia diffusion.