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.     

The relative uptake of Ca and Sr into tree foliage using a whole-watershed calcium addition

The use of strontium isotopes and ratios of alkaline earth elements (i.e., ⁸⁷Sr/⁸⁶Sr and Ca/Sr) to trace Ca sources to plants has become common in ecosystem studies. Here we examine the relative uptake of Ca and Sr in trees and subsequent accumulation in foliage. Using a whole-watershed Ca addition experiment at the Hubbard Brook Experimental Forest in N.H., we measured the uptake of Ca relative to Sr in foliage and roots of sugar maple (Acer saccharum), yellow birch (Betula alleghaniensis), American beech (Fagus grandifolia), and red spruce (Picea rubens). Vegetation was analyzed for Ca and Sr concentrations and the ⁸⁷Sr/⁸⁶Sr ratio. A comparison of the Ca/Sr ratio in the vegetation and the Ca/Sr ratio of the applied mineral allows for the calculation of a discrimination factor, which defines whether Ca and Sr are incorporated and allocated in the same ratio as that which is available. A discrimination factor greater than unity indicates preferential uptake of Ca over Sr; a factor less than unity reflects preferential uptake of Sr over Ca. We demonstrate that sugar maple (SM) and yellow birch (YB) have similar and small discrimination factors (1.14 ± 0.12,1σ and 1.16 ± 0.09,1σ) in foliage formation and discrimination factors of less than 1 in root formation (0.55–0.70). Uptake into beech suggests a larger discrimination factor (1.9 ± 1.2) in foliage but a similar root discrimination factor to SM and YB (0.66 ± 0.06,1σ). Incorporation into spruce foliage occurs at a much slower rate than in these other tree species and precludes evaluation of Ca and Sr discrimination in spruce foliage at this time. Understanding the degree to which Ca is fractionated from Sr in different species allows for refinement in the use of ⁸⁷Sr/⁸⁶Sr and Ca/Sr ratios to trace Ca sources to foliage. Methods from this study can be applied to natural environments in which various soil cation pools have different ⁸⁷Sr/⁸⁶Sr and Ca/Sr ratios. The results reported herein have implications for re-evaluating Ca sources and fluxes in forest ecosystems.     

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.     

Long-term records of strontium isotopic composition in tree rings suggest changes in forest calcium sources in the early 20th century

Many studies made in Europe and North America have shown an increasing depletion of exchangeable base cations that may cause tree nutritional deficiencies in sensitive soils. We use radial variation of strontium isotope in tree-rings (⁸⁷Sr/⁸⁶Sr ratio) to monitor possible changes in Ca sources for tree nutrition (Sr is used as an analog to Ca). The two main sources of Ca in forest stands are mineral weathering release and atmospheric inputs. Measurements in several forest stands in temperate regions show a steep decrease from pith to outer wood of the Sr isotope ratio from∼1870 to∼1920 except for stands developed on soils with a higher Ca status. This suggests a decrease of the weathering contribution (high ⁸⁷Sr/⁸⁶Sr ratio) when cations are displaced from the soil exchange complex by acid deposition at a rate faster than the replenishment of the cation pool by mineral weathering. This displacement enhances the atmospheric contribution, which is characterized by a low ⁸⁷Sr/⁸⁶Sr ratio. Tree-ring chronologies are an exceptional historic-timing record of chemical changes in the soil environment induced by atmospheric pollution. The reliability of the tree-ring recorder has been verified with a well-controlled nutritional perturbation in the context of a limed forest stand (with a known liming Sr isotopic signature). Our data suggest that forest ecosystems were affected by atmospheric inputs of strong acids earlier than previously thought.     

Determination of foliar Ca/Sr discrimination factors for six tree species and implications for Ca sources in northern hardwood forests

Plant and Soil - Discrimination during foliar uptake between the alkaline earth elements Ca and Sr must be understood to fully utilize Ca/Sr and 87Sr/86Sr ratios as a monitor of Ca sources to...     

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.     

Distinguishing sources of base cations in irrigated and natural soils: evidence from strontium isotopes

Strontium isotope ratios (⁸⁷Sr/⁸⁶Sr) of soil solids, soil cation extracts, irrigation water and plant material are used to determine strontium and therefore cation sources and fluxes in irrigated and natural soil–plant systems. Strontium isotopes of soil solids from four soil profiles (two irrigated vineyard soils and two 'natural' profiles from nearby reserves) show large differences between soil horizons with depth. These differences are not reflected in ⁸⁷Sr/⁸⁶Sr ratios of soil labile cations, which show both little variation down-profile and much lower ratios than soil solids. In the undisturbed, natural soil profiles, labile cation ⁸⁷Sr/⁸⁶Sr ratios are close to the ratio of modern seawater, indicating that solutes of marine origin from precipitation are the primary input of strontium (and calcium) to the labile cation pool. In the irrigated soil profiles, ⁸⁷Sr/⁸⁶Sr ratios of labile cations are consistently shifted towards that of the irrigation water. Mass-balance calculations using ⁸⁷Sr/⁸⁶Sr ratios of the different inputs to the labile cation pool indicate more than 90% of labile strontium is derived from precipitation solutes in unirrigated soils, and up to 44% from irrigation water solutes as an additional source in irrigated soils. The ⁸⁷Sr/⁸⁶Sr ratios of grapes grown in irrigated soils match precisely with those of the labile soil cations, demonstrating that cation nutrients are drawn wholly from the labile cation pool and have the same mix of precipitation, irrigation, and soil solid sources. The ⁸⁷Sr/⁸⁶Sr ratios of grapes grown in the irrigated soils may therefore vary over time depending on (1) the changing mix of irrigation water and local precipitation and (2) potential change to irrigation water. These findings suggest limitations to the use of strontium isotopes in the tracing of grapes and wines to their soil of origin.     

Localisation of mineral uptake by roots using Sr isotopes

To assess the contribution of deep soil horizons to the mineral supply of trees, we investigated the natural variation in the⁸⁷ Sr/⁸⁶Sr isotopic ratio of plant-available strontium with soil depth. In three sites of North-western Spain, this ratio increased with soil depth. The comparison of isotopic ratios of tree leaves and roots at different depths showed that most of the Sr accumulation in Eucalyptus globulus and Pinus pinaster growing on shallow and poor soils in this rainy climate originated from the upper soil layers. As Ca and Sr behave similarly in the soil-plant system, this conclusion can be applied to Ca. This superficial uptake is attributed to the low availability of Sr and Ca in the soil as well as to the shortness of the drought period as compared to the length of the growth period. This technique appears to offer a promising way of studying relative root distributions in soils and plant competition for nutrients.     

Use of strontium isotopes and foliar K content to estimate weathering of biotite induced by pine seedlings colonised by ectomycorrhizal fungi from two different soils

Pinus sylvestris seedlings, colonised by ectomycorrhizal (EM) fungi from either of two different soils (untreated forest soil and a limed soil from a clear cut area), were grown with or without biotite as a source of K. The biotite was naturally enriched in ⁸⁷Sr and the ratio of ⁸⁷Sr/ ⁸⁶Sr in the plant biomass was estimated and used as a marker for biotite weathering and compared to estimates of weathering based on foliar content of K. Different nutrient regimes were used to expose the seedlings to deficiencies of K with and without an application of nitrogen (NH₄NO₃) in excess of seedling demand. The seedlings were grown for 220 days and the elemental composition of the shoots were analysed at harvest. The EM colonisation was followed by analysing the concentration of ergosterol in the roots and the soils. Bacterial activity of the soil was estimated by the thymidine incorporation technique. The concentration of organic acids in the soil solution was measured in the soil in which seedlings colonised by EM fungi from the untreated forest soil were grown. It was found that seedlings colonised by EM fungi from untreated forest soil had taken up more K in treatments with biotite addition compared to seedlings colonised by EM fungi from the limed forest soil (p<0.05). Seedlings from untreated forest soil had larger shoots and contained more K when grown with biotite compared to KCl as K source, indicating that biotite had a stimulatory effect on the growth of these seedlings which was not related to K uptake. Seedlings from the limed soil, on the other hand, had similar foliar K content when grown with either biotite or KCl as K source. The larger uptake of K in seedlings from untreated forest soil was not an effect of a more developed EM colonisation of the roots since seedlings from the limed soil had a higher ergosterol concentration both in the soil and in the roots. Nutrient regimes had no significant influence on the total uptake of K but the ⁸⁷Sr/ ⁸⁶Sr isotope ratio in the plant biomass indicated that seedlings grown with excess nitrogen supply had taken up proportionally less Sr from the biotite (1.8% of total Sr content) compared to seedlings grown with a moderate nitrogen supply (5.0%). Furthermore, seedlings grown with excess nitrogen supply had a reduced fungal colonisation of roots and soil and bacterial activity was lower in these soils. The ⁸⁷Sr/ ⁸⁶Sr ratio in the plant biomass was positively correlated with fungal colonisation of the roots (r ²=0.98), which may indicate that the fungus was involved in releasing Sr from the biotite. Uptake of K from biotite was not related to the amount of organic acids in the soil solution. Oxalic acid was positively related to the amount of ergosterol in the root, suggesting that oxalic acid in the soil solution originates from the EM symbionts. The accuracy of the estimations of biotite weathering based on K uptake by the seedlings in comparison with the ⁸⁷Sr/⁸⁶Sr isotope ratio measured in the shoots is discussed. This revised version was published online in June 2006 with corrections to the Cover Date.     

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.     

Biomass, morphology and nutrient contents of fine roots in four Norway spruce stands

Fine root systems may respond to soil chemical conditions, but contrasting results have been obtained from field studies in non-manipulated forests with distinct soil chemical properties. We investigated biomass, necromass, live/dead ratios, morphology and nutrient concentrations of fine roots (<2 mm) in four mature Norway spruce (Picea abies [L.] Karst.) stands of south-east Germany, encompassing variations in soil chemical properties and climate. All stands were established on acidic soils (pH (CaCl₂) range 2.8–3.8 in the humus layer), two of the four stands had molar ratios in soil solution below 1 and one of the four stands had received a liming treatment 22 years before the study. Soil cores down to 40 cm mineral soil depth were taken in autumn and separated into four fractions: humus layer, 0–10 cm, 10–20 cm and 20–40 cm. We found no indications of negative effects of N availability on fine root properties despite large variations in inorganic N seepage fluxes (4–34 kg N ha⁻¹ yr⁻¹), suggesting that the variation in N deposition between 17 and 26 kg N ha⁻¹ yr⁻¹ does not affect the fine root system of Norway spruce. Fine root biomass was largest in the humus layer and increased with the amount of organic matter stored in the humus layer, indicating that the vertical pattern of fine roots is largely affected by the thickness of this horizon. Only two stands showed significant differences in fine root biomass of the mineral soil which can be explained by differences in soil chemical conditions. The stand with the lowest total biomass had the lowest Ca/Al ratio of 0.1 in seepage, however, Al, Ca, Mg and K concentrations of fine roots were not different among the stands. The Ca/Al ratio in seepage might be a less reliable stress parameter because another stand also had Ca/Al ratios in seepage far below the critical value of 1.0 without any signs of fine root damages. Large differences in the live/dead ratio were positively correlated with the Mn concentration of live fine roots from the mineral soil. This relationship was attributed to faster decay of dead fine roots because Mn is known as an essential element of lignin degrading enzymes. It is questionable if the live/dead ratio can be used as a vitality parameter of fine roots since both longevity of fine roots and decay of root litter may affect this parameter. Morphological properties were different in the humus layer of one stand that was limed in 1983, indicating that a single lime dose of 3–4 Mg ha⁻¹ has a long-lasting effect on fine root architecture of Norway spruce. Almost no differences were found in morphological properties in the mineral soil among the stands, but vertical patterns were apparently different. Two stands with high base saturation in the subsoil showed a vertical decrease in specific root length and specific root tip density whereas the other two stands showed an opposite pattern or no effect. Our results suggest that proliferation of fine roots increased with decreasing base saturation in the subsoil of Norway spruce stands.     

Does soil acidity reduce subsoil rooting in Norway spruce (Picea abies)?

Increasing evidence suggests that forest soils in central and northern Europe as well as in North America have been significantly acidified by acid deposition during the last decades. The present investigation was undertaken to examine the effect of soil acidity on rooting patterns of 40-year-old Norway spruce trees by comparing fine and coarse roots among four stands which differed in soil acidity and Mg (and Ca) nutrition. The coarse root systems of four to five 40-year-old Norway spruce trees per stand were manually excavated. The sum of cross sectional area (CSA) at 60 cm soil depth and below of all vertical coarse roots, as a measure of vertical rooting intensity, was strongly reduced with increasing subsoil acidity of the stands. This pattern was confirmed when 5 additional acidic sites were included in the analysis. Fine root biomass in the mineral soil estimated by repeated soil coring was strongly reduced in the heavily acidified stands, but increased in the humic layer. Using ingrowth cores and a screen technique, we showed that the higher root biomass in the humic layer of the more acidic stands was a result of higher root production. Thus, reduced fine root biomass and coarse root CSA in deeper soil layers coincided with increased root growth in the humic layer. Root mineral analysis showed Ca/Al ratios decreased with decreasing base saturation in the deeper mineral soil (20–40 cm). In the top mineral soil, only minor differences were observed among stands. In general, low Ca/Al ratios coincided with low fine root biomass. Calcium/aluminum ratios determined in cortical cell walls using X-ray microanalysis showed a similar pattern as Ca/Al ratios based on analysis of whole fine roots, although the amplitude of changes among the stands was much greater. Aluminum concentrations and Ca/Al ratios in cortical cell walls were at levels found to inhibit root growth of spruce seedlings in laboratory experiments. The data support the idea that Al (or Ca/Al ratios) and acid deposition-induced Mg (and possibly Ca) deficiency are important factors influencing root growth and distribution in acidic forest soils. Changes in carbon partitioning within the root system may contribute to a reduction in deep root growth.     

Wood-ash recycling affects forest soil and tree fine-root chemistry and reverses soil acidification

Wood ash was applied to a forest ecosystem with the aim to recycle nutrients taken from the forest and to mitigate the negative effects of intensive harvesting. After two years, the application of 8,000 kg ha⁻¹ of wood ash increased soil exchangeable Ca and Mg. Similarly, an increase in Ca and Mg in the Norway spruce fine roots was recorded, leading to significant linear correlations between soil and root Ca and soil and root Mg. In contrast to these macronutrients, the micronutrients Fe and Zn and the toxic element Al decreased in the soil exchangeable fraction with the addition of wood ash, but not in the fine roots. Only Mn decreased in soil and in fine roots leading to a significant linear correlation between soil and root Mn. In soil, as well as in fine roots, strong positive correlations were found between the elements Ca and Mg and between Fe and Al. This indicates that the uptake of Mg resembles that of Ca and that of Al that of Fe. With the wood ash application, the pH increased from 3.2 to 4.8, the base saturation from 30% to 86%, the molar basic cations/Al ratio (BC/Al) of the soil solution from 1.5 to 5.5, and the molar Ca/Al ratio of the fine roots from 1.3 to 3.7. Overall, all below-ground indicators of soil acidification responded positively to the wood ash application within two years. Nitrate concentrations increased only slightly in the soil solution at a soil depth of 75–80 cm, and no signs of increased heavy metal concentrations in the soils or in the fine roots were apparent. This suggests that the recycling of wood ash could be an integral part of sustainable forest management because it closes the nutrient cycle and reverses soil acidification.     

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.     

Dissolution of wollastonite during the experimental manipulation of Hubbard Brook Watershed 1

Powdered and pelletized wollastonite (CaSiO₃) was applied to an 11.8 ha forested watershed at the Hubbard Brook Experimental Forest (HBEF) in northern New Hampshire, U.S.A. during October of 1999. The dissolution of wollastonite was studied using watershed solute mass balances, and a ⁸⁷Sr/⁸⁶Sr isotopic tracer. The wollastonite (⁸⁷Sr/⁸⁶Sr = 0.70554) that was deposited directly into the stream channel began to dissolve immediately, resulting in marked increases in stream water Ca concentrations and decreases in the ⁸⁷Sr/⁸⁶Sr ratios from pre-application values of 0.872 mg/L and 0.72032 to values of 2.6 mg/L and 0.71818 respectively. After one calendar year, 401 kg of the initial 631 kg of wollastonite applied to the stream channel was exported as stream dissolved load, and 230 kg remained within the stream channel as residual CaSiO₃ and/or adsorbed on streambed exchange sites. Using previously established values for streambed Ca exchange capacity at the HBEF, the dissolution rate for wollastonite was found to be consistent with dissolution rates measured in laboratory experiments. Initially, Ca was released from the mineral lattice faster than Si, resulting in the development of a Ca-depleted leached layer on mineral grains. The degree of preferential Ca release decreased with time and reached stoichiometric proportions after 6 months. Using Sr as a proxy for Ca, the Ca from wollastonite dissolution can be accurately tracked as it is transported through the aquatic and terrestrial ecosystems of this watershed.     

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.     

Fine Root Production and Turnover in a Norway Spruce Stand in Northern Sweden: Effects of Nitrogen and Water Manipulation

Fine root length production, biomass production, and turnover in forest floor and mineral soil (0–30 cm) layers were studied in relation to irrigated (I) and irrigated-fertilized (IL) treatments in a Norway spruce stand in northern Sweden over a 2-year period. Fine roots (<1 mm) of both spruce and understory vegetation were studied. Minirhizotrons were used to estimate fine root length production and turnover, and soil cores were used to estimate standing biomass. Turnover was estimated as both the inverse of root longevity (RTL) and the ratio of annual root length production to observed root length (RTR). RTR values of spruce roots in the forest floor in I and IL plots were 0.6 and 0.5 y⁻¹, respectively, whereas the corresponding values for RTL were 0.8 and 0.9 y⁻¹. In mineral soil, corresponding values for I, IL, and control (C) plots were 1.2, 1.2, and 0.9 y⁻¹ (RTR) and 0.9, 1.1, and 1 y⁻¹ (RTL). RTR and RTL values of understory vegetation roots were 1 and 1.1 y⁻¹, respectively. Spruce root length production in both the forest floor and the mineral soil in I plots was higher than in IL plots. The IL-treated plots gave the highest estimates of spruce fine root biomass production in the forest floor, but, for the mineral soil, the estimates obtained for the I plots were the highest. The understory vegetation fine root production in the I and IL plots was similar for both the forest floor and the mineral soil and higher (for both layers) than in C plots. Nitrogen (N) turnover in the forest floor and mineral soil layers (summed) via spruce roots in IL, I, and C plots amounted to 2.4, 2.1, and 1.3 g N m⁻² y⁻¹, and the corresponding values for field vegetation roots were 0.6, 0.5, and 0.3 g N m⁻² y⁻¹. It was concluded that fertilization increases standing root biomass, root production, and N turnover of spruce roots in both the forest floor and mineral soil. Data on understory vegetation roots are required for estimating carbon budgets in model studies.     

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.     

Clean rain promotes fine root growth and soil respiration in a Norway spruce forest

Soil acidification and N saturation are considered to affect the decomposition of soil organic matter as well as growth and mortality of fine roots in many forest soils. Here we report from a field experiment where ‘clean rain’ has been applied to the soil for about 10 years under a roofed plot of a 71‐year‐old Norway spruce plantation at Solling, Central Germany. Reduced amounts of protons (?78%), sulphate (?53%), ammonium (?86%), and nitrate (?49%) were sprayed on the soil surface of the clean rain plot between 1992 and 2001. In an adjacent roofed control plot, throughfall was collected and immediately re‐sprinkled below the roof construction without any chemical manipulation. One year before the clean rain treatment started, live and dead fine root masses (≤2 mm) were determined from undisturbed soil cores down to 40 cm mineral soil depth. Total live fine root mass was significantly lower in the clean rain plot than in the control plot. After the first sampling, the soil holes were refilled with quartz sand and repeatedly sampled in June 1992, June 1996, and October 2001. There were no differences in live and dead fine root masses between the plots in 1992 and 1996. In 2001, both live and dead fine root masses of the clean rain plot were about twice as high as in the control plot, indicating that fine root growth recovered in the mineral soil following 10 years of clean rain treatment. Moreover, the clean rain treatment significantly reduced the total N concentrations of live fine roots and 1‐year‐old needles. Our results suggest that the reduced N input promoted fine root growth to compensate N deficiency. Reduced Al concentration in soil solution may have contributed to the recovery of fine root growth, however, the toxicity of Al species is largely unknown. Mean annual soil respiration rate was 24% higher in the period from 2000 to 2001, indicating that the clean rain treatment increased respiration of roots and heterotrophic microorganisms within the rhizosphere. Laboratory incubation of samples from the organic horizon and the top mineral soil revealed no differences between the plots in the decay rate of soil organic matter. Our results suggest that strong reductions in atmospheric N deposition from about 30 to 10 kg N ha^?1 yr^?1 and decreasing acid stress can have beneficial effects on growth of fine roots in the mineral soil within a decade. We conclude that biological recovery under reduced atmospheric loads can affect the nutrient and carbon budget of spruce soils in the long run.