Use of foliar Ca/Sr discrimination and <Superscript>87</Superscript>Sr/<Superscript>86</Superscript>Sr ratios to determine soil Ca sources to sugar maple foliage in a northern hardwood forest

Calcium/strontium and ⁸⁷Sr/⁸⁶Sr ratios in foliage can be used to determine the relative importance of different soil sources of Ca to vegetation, if the discrimination of Ca/Sr by the plant between nutrient sources and foliage is known. We compared these tracers in the foliage of sugar maple (Acer saccharum) to the exchange fraction and acid leaches of soil horizons at six study sites in the White Mountains of New Hampshire, USA. In a previous study, sugar maple was shown to discriminate for Ca compared to Sr in foliage formation by a factor of 1.14 ± 0.12. After accounting for the predicted 14% shift in Ca/Sr, foliar Ca/Sr and ⁸⁷Sr/⁸⁶Sr ratios closely match the values in the Oie horizon at each study site across a 3.6-fold variation in foliar Ca/Sr ratios. Newly weathered cations, for which the Ca/Sr and ⁸⁷Sr/⁸⁶Sr ratios are estimated from acid leaches of soils, can be ruled out as a major Ca source to current foliage. Within sites, the ⁸⁷Sr/⁸⁶Sr ratio of the soil exchange pool in the Oa horizon and in the 0–10 cm and 10–20 cm increments of the mineral soil are similar to the Oie horizon and sugar maple foliar values, suggesting a common source of Sr in all of the actively cycling pools, but providing no help in distinguishing among them as sources to foliage. The Ca/Sr ratio in the soil exchange pool, however, decreases significantly with depth, and based on this variation, the exchange pool below the forest floor can be excluded as a major Ca source to the current sugar maple foliage. This study confirms that internal recycling of Ca between litter, organic soil horizons and vegetation dominate annual uptake of Ca in northern hardwood ecosystems. Refinement of our understanding of Ca and Sr uptake and allocation in trees allows improvement in the use of Ca/Sr and ⁸⁷Sr/⁸⁶Sr ratios to trace Ca sources to plants.     

A comparison between Ca and Sr cycling in forest ecosystems

In favourable conditions, the ⁸⁷Sr/⁸⁶Sr isotope ratios of the Sr delivered by rain and soil mineral weathering differ. Assuming that Ca and Sr behave similarly in forest ecosystems, several authors have used the ⁸⁷Sr/⁸⁶Sr variation in forest compartments to calculate the contribution of rain and mineral weathering to Ca fluxes and pools. However, there are a number of experimental reports showing that Ca and Sr may behave differently in the soil and in the plant. We have tested this Ca–Sr analogy in the field by measuring the variation of Sr and Ca concentrations, fluxes and pools in spruce, beech and maple stands on granite, sandstone and limestone. Results show that (1) variations of Ca and Sr concentrations are generally correlated at each level of the ecosystems. (2) In spruce on acid soils, a preferential uptake of Ca over Sr occurs (Aubure spruce Sr/Ca = 0.8×10⁻³; soil exchangeable Sr/Ca between 2 and 6×10⁻³). On calcareous soils, a preferential uptake of Sr over Ca by spruce may occur. (3) In spruce and beech on acid and calcareous soils, a preferential translocation of Ca over Sr from roots to leaves occurs ((Sr/Ca) in leaves was between 10 and 90% of that in roots). (4) The biological cycling of Ca and Sr leads to an enrichment of the upper soil layers in Ca and Sr. Compared to Sr, Ca accumulates in the upper layer of acid soils because Ca cycling through litterfall is favoured over Sr cycling, and possibly because of the selectivity of acid organic exchangers for Ca. This revised version was published online in June 2006 with corrections to the Cover Date.     

Variations of bioavailable Sr concentration and 87Sr/86Sr ratio in boreal forest ecosystems

The mean depth of Sr and water uptake in mixed Norway spruce (Picea abies) and Scots pine (Pinus sylvestris) stands was investigated, using natural variations of ⁸⁷Sr/⁸⁶Sr and ¹⁸O/¹⁶O in soils in relation to depth. Three spruce-pine pairs were studied on a podzol and a peat site in Northern Sweden. Tree leaf and wood, as well as soils, soil solutions and roots below each tree were analysed for Sr and Ca concentrations and ⁸⁷Sr/⁸⁶Sr ratio. The ¹⁸O/¹⁶O ratio was also determined in xylem sap and soil solutions in relation to depth. Soil solution ¹⁸O/¹⁶O decreased in relation to depth. Comparing with xylem sap ¹⁸O/¹⁶O data indicated a deeper uptake of soil water by pine than spruce on the podzol site and a superficial uptake by both species on the peat. The ⁸⁷Sr/⁸⁶Sr ratio of bioavailable Sr generally increased in soils in relation to depth. Contrastingly, the ⁸⁷Sr/⁸⁶Sr ratio in spruce wood was generally higher than in pine wood suggesting a deeper uptake of Sr by spruce. But the ⁸⁷Sr/⁸⁶Sr ratio and concentrations of bioavailable Sr were systematically higher below spruce than below pine. In order to explain these unexpected results, we built a simple flux model to investigate the possible effects of interspecific variations in Sr cycling, soil mineral weathering and depth of Sr uptake on soil and tree ⁸⁷Sr/⁸⁶Sr ratio. At the study sites, spruce cycled in litterfall up to 12 times more strontium than pine. The use of the model showed that this difference in Sr cycling could alone explain higher isotopic signatures of trees and topsoils below spruce. Besides, high isotopic signatures of roots in the A/E horizons below spruce led us to hypothesise a species-specific weathering process. Finally, the comparison between the ⁸⁷Sr/⁸⁶Sr ratios in wood and root or soil solutions below each species suggested that the average depth of Sr and water uptake were close, but irregular variations of the Sr isotopic ratio with depth reduce the accuracy of the results. Tree species strongly influence Sr isotopic ratios in boreal forest soils through differences in Sr cycling, and possibly through specific mineral weathering.     

Test of soil extractants for their suitability in predicting Ca/Sr ratios in leaves and stems of sugar maple seedlings

Differential uptake and translocation of Ca and Sr in organisms have been reported, calling into question the use of Sr to track Ca cycling in the environment. We investigated the relationship between Ca/Sr ratios in soil extracts of various strengths (H₂O, NH₄Cl, and NH₄EDTA) and seedlings of sugar maple (Acer saccharum Marsh.) grown from natural regeneration on 37 sites. Our objectives were to determine if Ca/Sr ratios in soil extracts are correlated with those in sugar maple tissues, and what soil extractant best duplicate plant tissue Ca/Sr ratios. Leaves had higher Ca/Sr ratios than stems and the extractants did not produce equal Ca/Sr ratios: H₂O had the lowest Ca/Sr, and NH₄EDTA the highest. The relationships between soil extract Ca/Sr ratios and leaf and stem Ca/Sr ratios were significant and linear, but the slopes differed among extractants. The lowest slope (0.45) was observed for the water extract/leaves and the highest (2.15) for the NH₄EDTA extract/stem with discrimination factors ranging from 0.22 with NH₄EDTA to 1.59 for water. Leaf extracts were more strongly correlated with soil Ca/Sr than stem extracts (R ² of 0.57–0.7 vs. R ² of 0.45–0.6, respectively). These findings support the use of Ca/Sr ratios in plants to track their source of soil Ca, but they highlight the need to calibrate the relationships for the plant tissue and soil extractant used.     

Do deep tree roots provide nutrients to the tropical rainforest?

The contribution of deep tree roots to the nutrition of a tropicalrainforest were studied along an edaphic transect in French Guyana. Soil typeswere mapped in relation to the texture of the upper horizons and the depth ofoccurrence of the loamy saprolite. The position of mature individuals of fourcommon species, differing by they rooting depth, was identified and tree leaveswere analysed for major nutrients and strontium (Sr) isotopic ratios.On average, the range of leaf isotopic ratio (⁸⁷Sr/⁸⁶Sr= 0.714–0.716) was narrow compared to that of bulk soils(⁸⁷Sr/⁸⁶Sr = 0.72–0.77). Steep gradients ofincreasing ⁸⁷Sr/⁸⁶Sr in roots with soil depth were foundin all investigated profiles, which indicated that the flux of Sr deposited inrain and leached from the litter layer was tightly retained in the upper soillayers. Over the whole of the site, as well as within each soil unit, tree⁸⁷Sr/⁸⁶Sr ratios were very similar whatever the species,and close to litter and near-surface roots ⁸⁷Sr/⁸⁶Srratios, suggesting no or very little Sr contribution from deep tree roots.Variations of Ca and Sr concentrations in leaves were strongly correlated butnot with leaf ⁸⁷Sr/⁸⁶Sr ratios. These results support thetheory that Sr and Ca uptake and cycling are mostly superficial in tropicalrainforests.     

Identifying calcium sources at an acid deposition-impacted spruce forest: a strontium isotope,alkaline earth element multi-tracer approach

Depletion of calcium from forest soils has important implications for forest productivity and health. Ca is available to fine feeder roots from a number of soil organic and mineral sources, but identifying the primary source or changes of sources in response to environmental change is problematic. We used strontium isotope and alkaline earth element concentration ratios of trees and soils to discern the record of Ca sources for red spruce at a base-poor, acid deposition-impacted watershed. We measured ⁸⁷Sr/⁸⁶Sr and chemical compositions of cross-sectional stemwood cores of red spruce, other spruce tissues and sequential extracts of co-located soil samples. ⁸⁷Sr/⁸⁶Sr and Sr/Ba ratios together provide a tracer of alkaline earth element sources that distinguishes the plant-available fraction of the shallow organic soils from those of deeper organic and mineral soils. Ca/Sr ratios proved less diagnostic, due to within-tree processes that fractionate these elements from each other. Over the growth period from 1870 to 1960, ⁸⁷Sr/⁸⁶Sr and Sr/Ba ratios of stemwood samples became progressively more variable and on average trended toward values that considered together are characteristic of the uppermost forest floor. In detail the stemwood chemistry revealed an episode of simultaneous enhanced uptake of all alkaline earth elements during the growth period from 1930 to 1960, coincident with reported local and regional increases in atmospheric inputs of inorganic acidity. We attribute the temporal trends in stemwood chemistry to progressive shallowing of the effective depth of alkaline earth element uptake by fine roots over this growth period, due to preferential concentration of fine roots in the upper forest floor coupled with reduced nutrient uptake by roots in the lower organic and upper mineral soils in response to acid-induced aluminum toxicity. Although both increased atmospheric deposition and selective weathering of Ca-rich minerals such as apatite provide possible alternative explanations of aspects of the observed trends, the chemical buffering capacity of the forest floor-biomass pool limits their effectiveness as causal mechanisms.     

Strontium-calcium discriminationin vitro by rat tissues

Uptake of⁸⁹Sr and⁴⁵Ca by 15 soft tissues of adult rat was studiedin vitro to assess the extent of discrimination between Sr and Ca. While brain, kidney, placenta and uterus have lower uptake of⁸⁹Sr and 45 Ca that of diaphragm, lactating mammary gland, skeletal muscle, skin, spleen and testes is higher. Tissues with medium range uptake are heart, small intestine, liver, lung, non-lactating mammary gland and ovary. The 6 tissues displaying discriminating ability, as expressed by⁸⁹Sr/⁴⁵Ca (tissue/medium), in the decreasing order are: small intestine, kidney, lactating mammary gland, placenta, diaphragm and heart. Non-lactating mammary gland and the other tissues did not differentiate between Sr and Ca. The efect of several enzyme inhibitors, compounds influencing Sr-Ca metabolism and other factors was studied in terms of the nature and mechanism of Sr-Ca discrimination.     

Calcium weathering in forested soils and the effect of different tree species

Soil weathering can be an important mechanism to neutralize acidity inforest soils. Tree species may differ in their effect on or response to soilweathering. We used soil mineral data and the natural strontium isotope ratio⁸⁷Sr/⁸⁶Sr as a tracer to identify the effect of treespecies on the Ca weathering rate. The tree species studied were sugar maple(Acer saccharum), hemlock (TsugaCanadensis), American beech (Fagusgrandifolia),red maple (Acer rubrum), white ash (FraxinusAmericana) and red oak (Quercus rubra) growingin a forest in northwestern Connecticut, USA. Three replicated sites dominatedby one of the six tree species were selected. At sugar maple and hemlock sitesthe dominant mineral concentrations were determined at three soil depths. Ateach site soil, soil water and stem wood of the dominant tree species weresampled and analyzed for the ⁸⁷Sr/⁸⁶Sr ratio, total SrandCa content. Atmospheric deposition was collected and analyzed for the sameconstituents. Optical analysis showed that biotite and plagioclaseconcentrations were lower in the soil beneath hemlock than beneath sugar mapleand suggested species effects on mineral weathering in the upper 10cm of the mineral soil. These results could not be confirmed withdata obtained by the Sr isotope study. Within the sensitivity of the Sr isotopemethod, we could not detect tree species effects on Ca weathering andcalculatedCa weathering rates were low at all sites (< 60mgm^–2yr^–1). Wefound a positive correlation between Ca weathering and the total Caconcentration in the surface soil. These results indicate that the absolutedifferences in Ca weathering rate between tree species in these acidic surfacesoils are small and are more controlled by the soil parent material(plagioclasecontent) than by tree species.     

Natural 15N abundance of soil N pools and N2O reflect the nitrogen dynamics of forest soils

Natural ¹⁵N abundance measurements of ecosystem nitrogen (N) pools and ¹⁵N pool dilution assays of gross N transformation rates were applied to investigate the potential of δ¹⁵N signatures of soil N pools to reflect the dynamics in the forest soil N cycle. Intact soil cores were collected from pure spruce (Picea abies (L.) Karst.) and mixed spruce-beech (Fagus sylvatica L.) stands on stagnic gleysol in Austria. Soil δ¹⁵N values of both forest sites increased with depth to 50 cm, but then decreased below this zone. δ¹⁵N values of microbial biomass (mixed stand: 4.7 ± 0.8‰, spruce stand: 5.9 ± 0.9‰) and of dissolved organic N (DON; mixed stand: 5.3 ± 1.7‰, spruce stand: 2.6 ± 3.3‰) were not significantly different; these pools were most enriched in ¹⁵N of all soil N pools. Denitrification represented the main N₂O-producing process in the mixed forest stand as we detected a significant ¹⁵N enrichment of its substrate NO₃⁻ (3.6 ± 4.5‰) compared to NH₄⁺ (−4.6 ± 2.6‰) and its product N₂O (−11.8 ± 3.2‰). In a ¹⁵N-labelling experiment in the spruce stand, nitrification contributed more to N₂O production than denitrification. Moreover, in natural abundance measurements the NH₄⁺ pool was slightly ¹⁵N-enriched (−0.4 ± 2.0 ‰) compared to NO₃⁻ (−3.0 ± 0.6 ‰) and N₂O (−2.1 ± 1.1 ‰) in the spruce stand, indicating nitrification and denitrification operated in parallel to produce N₂O. The more positive δ¹⁵N values of N₂O in the spruce stand than in the mixed stand point to extensive microbial N₂O reduction in the spruce stand. Combining natural ¹⁵N abundance and ¹⁵N tracer experiments provided a more complete picture of soil N dynamics than possible with either measurement done separately.     

Effect of soil K, Ca and Mg saturation and endomycorrhization on growth and nutrient uptake of sugar maple seedlings

Nutrient imbalances of declining sugar maple (Acer saccharum Marsh.) stands in southeastern Quebec have been associated with high exchangeable Mg levels in soils relative to soil K and Ca. A greenhouse experiment was set up to test the hypothesis that the equilibrium between soil exchangeable K, Ca, and Mg ions influences the growth and nutrient status of sugar maple seedlings. Also tested was whether endomycorrhization can alter nutrient acquisition under various soil exchangeable basic cations ratios. Treatments consisted of seven ratios of soil exchangeable K, Ca, and Mg making up a total base saturation of 58%, and a soil inoculation treatment with the endomycorrhizal fungus Glomus versiforme (control and inoculated), in a complete factorial design. Sugar maple seedlings were grown for 3 months in the treated soils. Plant shoot elongation rate, dry biomass and nutrient concentrations in foliage were influenced by the various ratios of soil cations. The predicted plant biomass and foliar K concentration were highest at a soil Ca saturation of 38%, a soil K saturation of 12%, and a soil Mg saturation of 8%. Potassium concentration in foliage was dependent on the level of Ca and Mg saturation in the soil when soil K saturation was close to 12%. Foliar Ca and Mg levels were more dependent on their corresponding levels in soil than foliar K. Colonization by G. versiforme did not influence seedling growth and macronutrient uptake. The results confirm that growth and nutrition of sugar maple are negatively affected by imbalances in exchangeable basic cations in soils.     

Silicon Isotopic Fractionation by Banana (Musa spp.) Grown in a Continuous Nutrient Flow Device

The determination of the plant-induced Si-isotopic fractionation is a promising tool to better quantify their role in the continental Si cycle. Si-isotopic signatures of the different banana plant parts and Si source were measured, providing the isotopic fractionation factor between plant and source. Banana plantlets (Musa acuminata Colla, cv Grande Naine) were grown in hydroponics at variable Si supplies (0.08, 0.42, 0.83 and 1.66 mM Si). Si-isotopic compositions were determined on a multicollector plasma source mass spectrometer (MC-ICP-MS) operating in dry plasma mode. Results are expressed as δ²⁹Si relative to the NBS28 standard, with an average precision of ± 0.08‰ (±2σ_D). The fractionation factor ²⁹ε between bulk banana plantlets and source solution is −0.40 ± 0.11‰. This confirms that plants fractionate Si isotopes by depleting the source solution in ²⁸Si. The intra-plant fractionation Δ²⁹Si between roots and shoots amounts to −0.21 ± 0.08‰. Si-isotopic compositions of the various plant parts indicate that heavy isotopes discrimination occurs at three levels in the plant (at the root epidermis, for xylem loading and for xylem unloading). At each step, preferential crossing of light isotopes leaves a heavier solution, and produces a lighter solution. Si-isotopic fractionation processes are further discussed in relation with Si uptake and transport in plants. These findings have important implications on the study of continental Si cycle.     

Evaluation of the mobility and discrimination of Ca, Sr and Ba in forest ecosystems: consequence on the use of alkaline-earth element ratios as tracers of Ca

A comprehensive understanding of Ca cycling in an ecosystem is desirable because of the role of this element in tree mineral nutrition and its status as a major base cation on the soil exchange complex. The determination of the origin of Ca in forests is particularly indicated in regard of important changes linked to acid inputs and intensive logging. Natural strontium isotopes are increasingly used as tracers of Ca in forest ecosystems for qualitative and quantitative assessments. Nevertheless this method is limited to relatively simple systems with two sources of nutrients. Some recent studies coupled Sr/Ca or Sr/Ba ratios to Sr isotopic measurements in order to solve more complex systems. Such method has however associated with it some uncertainties: this approach assumed that Ca, Sr and Ba behave similarly throughout the ecosystem and does not take into account the Ca biopurification processes occurring in some tree’s organs which can alter element ratio. The present work focuses on two deciduous species covering large areas in Europe: European beech (Fagus sylvatica L.) and pedunculate oak (Quercus robur L.). In order to test the similarity of behaviour between Ca, Sr and Ba, their concentrations were measured extensively in the major compartments of two forest ecosystems. In parallel, the discrimination process inside tree organs was studied in 23 stands for beech and 10 stands for oak. We found that Sr and Ca behave similarly in all soil and tree compartments. By contrast, Ba and Ca appear to have contrasting behaviours, especially in streams, soil solution and soil exchange complex (no correlations between element concentrations). Sr/Ba and Ba/Ca ratios must therefore be used with care as tracer of Ca. The Ca biopurification is absent in roots and slight in bole wood but is large in bark, twigs and leaves. The discrimination factors (DF) between wood and leaves are characteristic of the two species studied and do not change significantly as a function of the soil Ca status (acidic or calcareous soils). Therefore, strontium–calcium DF can be used as a correction factor of the Sr/Ca ratio of leaves when this ratio is used in connection with Sr isotopic ratios. This correction allows to solve systems of tree nutrition with more than two sources of Ca.     

Asteriscus v. lapillus: comparing the chemistry of two otolith types and their ability to delineate riverine populations of common carp Cyprinus carpio

The chemical composition of common carp Cyprinus carpio asteriscus (vaterite) and lapillus (aragonite) otoliths from the same individual and reflecting the same growth period was measured to (1) determine whether there are differences in the uptake of trace metals (Mg:Ca, Mn:Ca, Sr:Ca and Ba:Ca ) and Sr isotope ratios (⁸⁷Sr:⁸⁶Sr) in co‐precipitating lapilli and asterisci and (2) compare the ability of multi‐element and isotopic signatures from lapilli, asterisci and both otolith types combined to discriminate C. carpio populations over a large spatial scale within a river basin. Depth profile analyses at the otolith edge using laser‐ablation inductively coupled plasma mass spectrometry showed that asterisci were enriched in Mg and Mn and depleted in Sr and Ba relative to lapilli, whilst ⁸⁷Sr:⁸⁶Sr values were nearly identical in both otolith types. Significant spatial differences among capture locations were found when all trace element and Sr isotope ratio data were aggregated into a multi‐element and isotopic signature, regardless of which otolith type was used or if they were used in combination. Discriminatory power was enhanced, however, when data for both otolith types were combined, suggesting that analysis of multiple otolith types may be useful for studies attempting to delineate C. carpio populations at finer spatial or temporal scales.     

Chronic Atmospheric NO<Stack><Subscript>3</Subscript><Superscript>−</Superscript></Stack> Deposition Does Not Induce NO<Stack><Subscript>3</Subscript><Superscript>−</Superscript></Stack> Use by <Emphasis Type="Italic">Acer saccharum</Emphasis> Marsh.

The ability of an ecosystem to retain anthropogenic nitrogen (N) deposition is dependent upon plant and soil sinks for N, the strengths of which may be altered by chronic atmospheric N deposition. Sugar maple (Acer saccharum Marsh.), the dominant overstory tree in northern hardwood forests of the Lake States region, has a limited capacity to take up and assimilate NO₃⁻. However, it is uncertain whether long-term exposure to NO₃⁻ deposition might induce NO₃⁻ uptake by this ecologically important overstory tree. Here, we investigate whether 10 years of experimental NO₃⁻ deposition (30 kg N ha⁻¹ y⁻¹) could induce NO₃⁻ uptake and assimilation in overstory sugar maple (approximately 90 years old), which would enable this species to function as a direct sink for atmospheric NO₃⁻ deposition. Kinetic parameters for NH₄⁺ and NO₃⁻ uptake in fine roots, as well as leaf and root NO₃⁻ reductase activity, were measured under conditions of ambient and experimental NO₃⁻ deposition in four sugar maple-dominated stands spanning the geographic distribution of northern hardwood forests in the Upper Lake States. Chronic NO₃⁻ deposition did not alter the V _max or K _m for NO₃⁻ and NH₄⁺ uptake nor did it influence NO₃⁻ reductase activity in leaves and fine roots. Moreover, the mean V _max for NH₄⁺ uptake (5.15 μmol ¹⁵N g⁻¹ h⁻¹) was eight times greater than the V _max for NO₃⁻ uptake (0.63 μmol ¹⁵N g⁻¹ h⁻¹), indicating a much greater physiological capacity for NH₄⁺ uptake in this species. Additionally, NO₃⁻ reductase activity was lower than most values for woody plants previously reported in the literature, further indicating a low physiological potential for NO₃⁻ assimilation in sugar maple. Our results demonstrate that chronic NO₃⁻ deposition has not induced the physiological capacity for NO₃⁻ uptake and assimilation by sugar maple, making this dominant species an unlikely direct sink for anthropogenic NO₃⁻ deposition.     

Estimating the relative nutrient uptake from different soil depths in Quercus robur, Fagus sylvatica and Picea abies

The distribution of fine roots and external ectomycorrhizal mycelium of three species of trees was determined down to a soil depth of 55 cm to estimate the relative nutrient uptake capacity of the trees from different soil layers. In addition, a root bioassay was performed to estimate the nutrient uptake capacity of Rb⁺ and NH₄⁺ by these fine roots under standardized conditions in the laboratory. The study was performed in monocultures of oak (Quercus robur L.), European beech (Fagus sylvatica L.) and Norway spruce [Picea abies (L.) Karst.] on sandy soil in a tree species trial in Denmark. The distribution of spruce roots was found to be more concentrated to the top layer (0–11 cm) than that of oak and beech roots, and the amount of external ectomycorrhizal mycelia was correlated to the distribution of the roots. The uptake rate of [⁸⁶Rb⁺] by oak roots declined with soil depth, while that of beech or spruce roots was not influenced by soil depth. In modelling the nutrient sustainability of forest soils, the utilization of nutrient resources in deep soil layers has been found to be a key factor. The present study shows that the more shallow-rooted spruce can have a similar capacity to take up nutrients from deeper soil layers than the more deeply rooted oak. The distribution of roots and mycelia may therefore not be a reliable parameter for describing nutrient uptake capacity by tree roots at different soil depths.     

Changes in Sr/Ca, Ba/Ca and 87Sr/86Sr ratios between trophic levels in two forest ecosystems in the northeastern U.S.A.

The variability and biologicalfractionation of Sr/Ca, Ba/Ca and ⁸⁷Sr/⁸⁶Srratios were studied in a soil–plant–invertebrate–bird food chain in two forested ecosystems withcontrasting calcium availability in the northeasternU.S.A. Chemical measurements were made of the soilexchange pool, leaves, caterpillars, snails, and boththe femurs and eggshells of breeding insectivorousmigratory songbirds. ⁸⁷Sr/⁸⁶Sr values weretransferred up the food chain from the soil exchangepool to leaves, caterpillars, snails and eggshellswithout modification. Adult birds were the oneexception; their ⁸⁷Sr/⁸⁶Sr values generallyreflected those of lower trophic levels at each site,but were lower and more variable, probably becausetheir strontium was derived in part from foods intropical winter habitats where lower⁸⁷Sr/⁸⁶Sr ratios are likely to predominate. Sr/Ca and Ba/Ca ratios decreased at each successive trophiclevel, supporting previous suggestions that Sr/Ca andBa/Ca ratios can be used to identify the trophic levelat which an organism is primarily feeding. The changesin Sr/Ca and Ba/Ca ratios we measured for vegetationand insects were comparable to similar measurementsmade previously (but based on single samples of eachorganism) in an alpine ecosystem. Changes in Sr/Ca andBa/Ca ratios between birds and their food have notpreviously been measured, but the values we obtainedwere similar to those for herbivorous and carnivorousmammals at similar trophic levels. Our results provideevidence that supports the use of Sr/Ca ratios in thedetermination of human paleodiets and suggests thatSr/Ca ratios may also provide a useful tool in studiesof modern food webs. Furthermore, our findings suggestthat ⁹⁰Sr from nuclear fallout will notbioaccumulate in forests and that changes in Sr/Caratios between trophic levels will need to beconsidered in some cases when using⁸⁷Sr/⁸⁶Sr as a tracer of calciumbiogeochemistry.     

Estimates of biomass and primary productivity in a high-altitude maple forest of the west central Himalayas

The paper describes the biomass and productivity of maple (Acer cappadocicum) forest occurring at an altitude of 2,750 m in the west central Himalayas. Total vegetation biomass was 308.3 t ha⁻¹, of which the tree layer contributed the most, followed by herbs and shrubs. The seasonal forest-floor litter mass varied between 5.4 t ha⁻¹ (in rainy season) and 6.6 t ha⁻¹ (in winter season). The annual litter fall was 6.2 t ha⁻¹, of which leaf litter contributed the largest part (59% of the total litter fall). Net primary productivity of total vegetation was 19.5 t ha⁻¹ year⁻¹. The production efficiency of leaves (net primary productivity/leaf mass) was markedly higher (2.9 g g⁻¹ foliage mass year⁻¹) than those of the low-altitude forests of the region.     

Spatial and temporal variations in otolith chemistry and relationships with water chemistry: a useful tool to distinguish Atlantic salmon Salmo salar parr from different natal streams

Otolith elemental (Sr:Ca, Ba:Ca, Mn:Ca, Mg:Ca and Rb:Ca) and isotopic (⁸⁷Sr:⁸⁶Sr) profiles from several annual cohorts of juvenile Atlantic salmon Salmo salar were related to the physico‐chemical characteristics (chemical signatures, flow rate, temperature and conductivity) of their natal rivers over an annual hydrological cycle. Only Sr:Ca, Ba:Ca and ⁸⁷Sr:⁸⁶Sr in otoliths were determined by their respective ratios in the ambient water. Sr:Ca ratios in stream waters fluctuated strongly on a seasonal basis, but these fluctuations, mainly driven by water flow regimes, were not recorded in the otoliths. Otolith Sr:Ca ratios remained constant during freshwater residency at a given site and were exclusively related to water Sr:Ca ratios during low flow periods. While interannual differences in otolith elemental composition among rivers were observed, this variability was minor compared to geographic variability and did not limit classification of juveniles to their natal stream. Success in discriminating fish from different sites was greatest using Sr isotopes as it remained relatively constant across years at a given location.     

Monitoring uptake and contents of Mg,Ca and K in Norway spruce as influenced by pH and Al,using microprobe analysis and stable isotope labelling

In a model system using intact spruce trees (Picea abies [L.] Karst.) we followed the path of magnesium, calcium and potassium during uptake into the root and during long-range transport into the shoot, by multiple stable isotope labelling. The roots of two- and three-year-old spruce trees originating from soil culture were removed from the soil and, in part or in toto, exposed to labelling solutions containing the stable isotopes ²⁵Mg or ²⁶Mg, ⁴¹K and ⁴²Ca or ⁴⁴Ca. Optical-emission-spectroscopy (ICP-OES) of plant fractions and labelling solutions was combined with the quantitative analysis of stable isotope ratios in sections of shock frozen, cryosubstituted material using the laser-microprobe-mass-analyser (LAMMA). This combination allowed us to distinguish, both in bulk samples and on the cellular level between (i) the fraction of elements originally present in the plant before the start of the labelling, (ii) the material taken up from the labelling solution into the plant and (iii) any material released by the plant into the labelling solution.In single-root labelling experiments, roots of three-year-old spruce trees, grown in nursery soil, were exposed to various pH conditions. The exchange of Mg and Ca with the labelling solution was nearly 100% in the cell walls of the mycorrhized finest roots. This exchange was only slightly affected by a step down to pH 3.5. The absolute Mg and Ca content in the cell walls was moderately reduced by incubation at pH 3.5 and strongly reduced in the presence of Al at this pH. After a pH 3.5 and 2 mM Al treatment we found Al in the xylem cell walls and the cortex cell lumina at elevated concentrations. To analyse the combined effect of high Al and high proton concentrations on the long-range transport, we used a split-root system. The root mass of an intact two-year-old spruce tree, grown in mineral soil, was divided into even parts and both halves incubated in solutions with two sets of different stable isotopes of Mg and Ca (side A: no Al, ²⁵Mg and ⁴²Ca; side B: +Al, ²⁶Mg and ⁴⁴Ca) and ⁴¹K on both sides. We observed a large uptake of Mg, Ca and K into the plant and a pronounced release. The net uptake of all three elements was lower from the Al-doted solution. In cross-sections of the apical shoot we found after seven-day labelling period about 60–70% of the Mg and Ca and 30% of the K content in the xylem cell walls originating from both labelling solutions. The clear majority of the Mg and Ca label originated from the Al-doted side.     

Norway Maple (Acer platanoides) Invasion of a Natural Forest Stand: Understory Consequence and Regeneration Pattern

Norway maple (Acer platanoidesis) is invasive in a natural stand in suburban Ithaca, NY. To determine the understory pattern and consequences of a Norway maple invasion, I compared density and species richness under Norway maples and sugar maples (Acer saccharum). Mean sapling density was significantly lower (P<0.0027) under Norway maples (3.64/100 m²±1.6 SE) than under sugar maples (19.4/100 m²±4.4 SE). Mean sapling species richness was significantly lower (P<0.0018) under Norway maples (0.7/32 m²±0.18 SE) than under sugar maples (2.6/32 m²±0.48 SE). Likewise, Norway maple regeneration is more frequent under sugar maples than sugar maple regeneration: 57% of sugar maple plots had Norway maple saplings while 0% of Norway maple plots had sugar maple saplings. Two significant plot effects were found for presence–absence: Norway maple saplings grow under Norway maples with a significantly lower frequency (P<0.03) than under sugar maples; sugar maple saplings grow under Norway maples with a significantly lower frequency (P<0.000) than under sugar maples. Across the site, Norway maple saplings were the most abundant (29 saplings for 480 m²). The success of Norway maple regeneration and the reductions in total stem density beneath Norway maples is most likely the result of its strong competitive abilities, notably its high shade tolerance and abundant seed crops.