The mobility of nitrogen across tree-rings of Norway spruce (Picea abies L.) and the effect of extraction method on tree-ring δ¹5N and δ¹³C values.

RATIONALE: The use of stable nitrogen (N) isotope ratios (δ(15)N values) in dendroecological studies is often preceded by an extraction procedure using organic solvents to remove mobile N compounds from tree-rings. Although these mobile N compounds may be capable of distorting potential environmental signals in the tree-ring δ(15)N values, recent investigations question the necessity of such an extraction. METHODS: We used an on-going experiment with simulated elevated N deposition previously labelled with (15)N, in conjunction with control trees, to investigate the necessity of extracting mobile N compounds (using a rapid extraction procedure) for tree-ring δ(15)N and δ(13)C studies, as well as N and C concentration analyses. In addition, we examined the magnitude of radial redistribution of N across tree-rings of Norway spruce (Picea abies). RESULTS: The (15)N label, applied in 1995/96, was found in tree-rings as far back as 1951, although the increased N availability did not cause any significant relative increase in tree growth. The rapid extraction procedure had no significant effect on tree-ring δ(15)N or δ(13)C values in either labelled or control trees, or on N concentration. The C concentrations, however, were significantly higher after extraction in control samples, with the opposite effect observed in labelled samples. CONCLUSIONS: Our results indicate that the extraction of mobile N compounds through the rapid extraction procedure is not necessary prior to the determination of Norway spruce δ(15)N or δ(13)C values in dendrochemical studies. δ(15)N values, however, must be interpreted with great care, particularly when used as a proxy for the N status of trees, due to the very high mobility of N within the tree stem sapwood of Norway spruce over several decades.

Nitrogen dynamics in oak model ecosystems subjected to air warming and drought on two different soils.

Being tolerant to heat and drought, oaks are promising candidates for future forestry in view of climate change in Central Europe. Air warming is expected to increase, and drought decrease soil N availability and thus N supply to trees. Here, we conducted a model ecosystem experiment, in which mixed stands of young oaks (Quercus robur, Q. petraea and Q. pubescens) were grown on two different soils and subjected to four climate treatments during three growing seasons: air warming by 1-2 °C, drought periods (average precipitation reduction of 43-60%), a combination of these two treatments, and a control. In contrast to our hypotheses, neither air warming nor drought significantly affected N availability, whereas total amounts, vertical distribution and availability of soil N showed substantial differences between the two soils. While air warming had no effect on tree growth and N accumulation, the drought treatment reduced tree growth and increased, or tended to increase, N accumulation in the reduced biomass, indicating that growth was not limited by N. Furthermore, (15) N-labelling revealed that this accumulation was associated with an increased uptake of nitrate. On the basis of our results, climate change effects on N dynamics are expected to be less important in oak stands than reduced soil water availability.

Soil nitrogen dynamics in a river floodplain mosaic.

In their natural state, river floodplains are heterogeneous and dynamic ecosystems that may retain and remove large quantities of nitrogen from surface waters. We compared the soil nitrogen dynamics in different types of habitat patches in a restored and a channelized section of a Thur River floodplain (northeast Switzerland). Our objective was to relate the spatiotemporal variability of selected nitrogen pools (ammonium, nitrate, microbial nitrogen), nitrogen transformations (mineralization, nitrification, denitrification), and gaseous nitrogen emission (NO) to soil properties and hydrological processes. Our study showed that soil water content and carbon availability, which depend on sedimentation and inundation dynamics, were the key factors controlling nitrogen pools and processes. High nitrogen turnover rates were measured on gravel bars, characterized by both frequent inundation and high sediment deposition rates, as well as in low-lying alluvial forest patches with a fine-textured, nutrient-rich soil where anaerobic microsites probably facilitated coupled nitrification-denitrification. In contrast, soils of the embankment in the channelized section had comparatively small inorganic nitrogen pools and low transformation rates, particularly those related to nitrate production. Environmental heterogeneity, characteristic of the restored section, favors nitrogen removal by creating sites of high sedimentation and denitrification. Of concern, however, are the locally high NO efflux and the possibility that nitrate could leach from nitrification hotspots.

Sinks for nitrogen inputs in terrestrial ecosystems: a meta-analysis of 15N tracer field studies.

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.

Water content and bark thickness of Norway spruce (Picea abies) stems: phloem water capacitance and xylem sap flow.

To determine the relationship between phloem transport and changes in phloem water content, we measured temporal and spatial variations in water content and sucrose, glucose and fructose concentrations in phloem samples and phloem exudates of 70- and 30-year-old Norway spruce trees (Picea abies (L.) Karst.). Large temporal and spatial variations in phloem water content (1.4-2.6 mg mg(dw)(-1)) and phloem total sugar concentration (31-70 mg g(dw)(-1)) paralleled each other (r(2) = 0.83, P < 0.0001 for the temporal profile and r(2) = 0.96, P < 0.008 for the spatial profile), indicating that phloem water content depends on the total amount of sugar to be transferred. Changes in phloem water content were unrelated to changes in bark thickness. Maximum changes in phloem water content calculated from dendrometer readings were only 8-11% of the maximum measured changes in phloem water content, indicating that reversible changes in bark thickness did not reflect changes in internal water relations. We also studied the relationship between xylem sap velocity and changes in bark thickness in 70-year-old trees during summer 1999 and winter 1999-2000. Sap flow occurred sporadically throughout the winter, but there was no relationship between bark shrinkage or swelling and sap velocity. In winter, mean daily xylem sap velocity was significantly correlated with mean daily vapor pressure deficit and air temperature (P < 0.0001, in both cases). Changes in bark thickness corresponded with both short- and long-term changes in relative humidity, in both winter and summer. Under controlled conditions at > 0 degrees C, changes in relative humidity alone caused changes in thickness of boiled bark samples. Because living bark of Norway spruce trees contains large areas with crushed and dead sieve cell zones-up to 24% of the bark is air-filled space-we suggest that this space can compensate for volume changes in living phloem cells independently of total tissue water content. We conclude that changes in bark thickness are not indicative of changes in either phloem water capacitance or xylem sap flow.

The accumulation of the rare earth elements and of scandium in successive needle age classes of Norway spruce.

The endogenous concentrations of Sc, La, Ce, Sm, Eu, Tb, Yb, and Lu were determined by neutron activation analysis in up to five successive needle age classes of Norway spruce (Picea abies). Trees from nine sites over different bedrocks were sampled individually. Concentration values found are generally much lower than those reported in the literature. This is attributed to the careful removal of any aerosols or soil particles from the needle surface prior to analysis. The concentration of each element increases linearly with the needle age class, i.e., the accumulation can be characterized by just one parameter, the yearly increment. This pattern is followed at small as well as at large concentrations. The accumulation behavior of the investigated elements is identical to that of Si. The relative concentrations of the rare earth elements (REE) in the needles are similar to those in the earth crust. There are significant correlations between the individual REE and between Sc and La.

Rubidium and cesium in spruce needles. Concentrations and biodynamics.

The endogenous concentrations of Rb and Cs have been determined in needles of 56 trees (Norway spruce, Picea abies) from eight different sites. Analysis was done by instrumental neutron activation. Concentrations were found to have a very large range (3-28,000 ng Cs/g and 1-190 micrograms Rb/g). The values on a given site have a tendency toward a log-normal distribution. There is a significant correlation between Rb and Cs, but the correlation is not linear. The concentrations of both elements are a function of the needle age. They decrease smoothly, approaching a constant value, when going from needle age class 1 to 5. It is shown that one algebraic function describes this biodynamic behavior on all sites and at all concentration levels. The function and its parameters are discussed.