Isotope ratios and concentrations of sulfur and nitrogen in needles and soils of Picea abies stands as influenced by atmospheric deposition of sulfur and nitrogen compounds

Concentrations and natural isotope abundance of total sulfur and nitrogen as well as sulfate and nitrate concentrations were measured in needles of different age classes and in soil samples of different horizons from a healthy and a declining Norway spruce (Picea abies (L.) Karst.) forest in the Fichtelgebirge (NE Bavaria, Germany), in order to study the fate of atmospheric depositions of sulfur and nitrogen compounds. The mean δ¹⁵N of the needles ranged between −3.7 and −2.1 ‰ and for δ³⁴S a range between −0.4 and +0.9 ‰ was observed. δ³⁴S and sulfur concentrations in the needles of both stands increased continuously with needle age and thus, were closely correlated. The δ¹⁵N values of the needles showed an initial decrease followed by an increase with needle age. The healthy stand showed more negative δ¹⁵N values in old needles than the declining stand. Nitrogen concentrations decreased with needle age. For soil samples at both sites the mean δ¹⁵N and δ³⁴S values increased from −3 ‰ (δ¹⁵N) or +0.9 ‰ (δ³⁴S) in the uppermost organic layer to about +4 ‰ (δ¹⁵N) or +4.5 ‰ (δ³⁴S) in the mineral soil. This depth-dependent increase in abundance of ¹⁵N and ³⁴S was accompanied by a decrease in total nitrogen and sulfur concentrations in the soil. δ¹⁵N values and nitrogen concentrations were closely correlated (slope −0.0061 ‰ δ¹⁵N per μmol eq N g_dw ⁻¹), and δ³⁴S values were linearly correlated with sulfur concentrations (slope −0.0576 ‰ δ³⁴S per μmol eq S g_dw ⁻¹). It follows that in the same soil samples sulfur concentrations were linearly correlated with the nitrogen concentrations (slope 0.0527), and δ³⁴S values were linearly correlated with δ¹⁵N values (slope 0.459). A correlation of the sulfur and nitrogen isotope abundances on a Δ basis (which considers the different relative frequencies of ¹⁵N and ³⁴S), however, revealed an isotope fractionation that was higher by a factor of 5 for sulfur than for nitrogen (slope 5.292). These correlations indicate a long term synchronous mineralization of organic nitrogen and sulfur compounds in the soil accompanied by element-specific isotope fractionations. Based on different sulfur isotope abundance of the soil (δ³⁴S=0.9 ‰ for total sulfur of the organic layer was assumed to be equivalent to about −1.0 ‰ for soil sulfate) and of the atmospheric SO₂ deposition (δ³⁴S=2.0 ‰ at the healthy site and 2.3 ‰ at the declining site) the contribution of atmospheric SO₂ to total sulfur of the needles was estimated. This contribution increased from about 20 % in current-year needles to more than 50 % in 3-year-old needles. The proportion of sulfur from atmospheric deposition was equivalent to the age dependent sulfate accumulation in the needles. In contrast to the accumulation of atmospheric sulfur compounds nitrogen compounds from atmospheric deposition were metabolized and were used for growth. The implications of both responses to atmospheric deposition are discussed.     

Effects of land use on 15N natural abundance of soils in Ethiopian highlands

We used the natural abundance of ¹⁵N in soils in forests, pastures and cultivated lands in the Menagesha and Wendo-Genet areas of Ethiopia to make inferences about the N cycles in these ecosystems. Since we have described the history of these sites based on variations in ¹³C natural abundance, patterns of δ¹⁵N and δ¹³C values were compared to determine if shifts of ¹⁵N correlate with shifts of vegetation. At Menagesha, a > 500-yr-old planted forest, we found δ¹⁵N values from −8.8 to +3.5‰ in litter, from −3.5 to +4.5‰ in 0–10 cm soil layer, and from −1.5 to +6.8‰ at >20 cm soil depth. The low δ¹⁵N in litter and surface mineral soils suggests that a closed N cycle has operated for a long time. At this site, the low δ¹³C of the surface horizon and the high δ¹³C of the lower soil horizons is clear evidence of a long phase of C₄ grass dominance or cultivation of C₄ crops before the establishment of the forest >500 years ago. In contrast, at Wendo-Genet, high δ¹³C of soils reveals that most of the land has been uncovered by forests until recently. Soil δ¹⁵N was high throughout (3.4–9.8‰), and there were no major differences between forested, cultivated and pasture soils in δ¹⁵N values of surface mineral soils. The high δ¹⁵N values suggest that open N cycles operate in the Wendo-Genet area. From the points of view of soil fertility management, it is interesting that tall forest ecosystems with relatively closed N cycling could be established on the fairly steep slopes at Menagesha after a long period of grass vegetation cover or cultivation. This revised version was published online in June 2006 with corrections to the Cover Date.     

Characteristics of DOC Exported from Northern Hardwood Forests Receiving Chronic Experimental NO<Stack><Subscript>3</Subscript><Superscript>−</Superscript></Stack> Deposition

Abstract Sugar maple (Acer saccharum Marsh.)-dominated northern hardwood forests of the Great Lakes Region commonly receive elevated levels of atmospheric nitrate (NO₃⁻) deposition, which can alter belowground carbon (C) cycling. Past research has demonstrated that chronic experimental NO₃⁻ deposition (3 g N m⁻² y⁻¹ above ambient) elicits a threefold increase in the leaching loss of dissolved organic carbon (DOC). Here, we used DOC collected from tension-cup lysimeters to test whether increased DOC export under experimental NO₃⁻ deposition originated from forest floor or mineral soil organic matter (SOM). We used DOC radiocarbon dating to quantify C sources and colorimetric assays to measure DOC aromaticity and soluble polyphenolic content. Our results demonstrated that DOC exports are primarily derived from new C (<50-years-old) in the forest floor under both ambient and experimental NO₃⁻ deposition. Experimental NO₃⁻ deposition increased soluble polyphenolic content from 25.03 ± 4.26 to 49.19 ± 4.23 μg phenolic C mg DOC⁻¹, and increased total aromatic content as measured by specific UV absorbance. However, increased aromatic compounds represented a small fraction (<10%) of the total observed increased DOC leaching. In combination, these findings suggest that experimental NO₃⁻ deposition has altered the production or retention as well as phenolic content of DOC formed in forest floor, however exact mechanisms are uncertain. Further elucidation of the mechanism(s) controlling enhanced DOC leaching is important for understanding long-term responses of Great Lakes forests to anthropogenic N deposition and the consequences of those responses for aquatic ecosystems.     

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.     

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.     

Growth of ectomycorrhizal mycelia and composition of soil microbial communities in oak forest soils along a nitrogen deposition gradient

Deciduous forests may respond differently from coniferous forests to the anthropogenic deposition of nitrogen (N). Since fungi, especially ectomycorrhizal (EM) fungi, are known to be negatively affected by N deposition, the effects of N deposition on the soil microbial community, total fungal biomass and mycelial growth of EM fungi were studied in oak-dominated deciduous forests along a nitrogen deposition gradient in southern Sweden. In-growth mesh bags were used to estimate the production of mycelia by EM fungi in 19 oak stands in the N deposition gradient, and the results were compared with nitrate leaching data obtained previously. Soil samples from 154 oak forest sites were analysed regarding the content of phospholipid fatty acids (PLFAs). Thirty PLFAs associated with microbes were analysed and the PLFA 18:2ω6,9 was used as an indicator to estimate the total fungal biomass. Higher N deposition (20 kg N ha⁻¹ y⁻¹ compared with 10 kg N ha⁻¹ y⁻¹) tended to reduce EM mycelial growth. The total soil fungal biomass was not affected by N deposition or soil pH, while the PLFA 16:1ω5, a biomarker for arbuscular mycorrhizal (AM) fungi, was negatively affected by N deposition, but also positively correlated to soil pH. Other PLFAs positively affected by soil pH were, e.g., i14:0, a15:0, 16:1ω9, a17:0 and 18:1ω7, while some were negatively affected by pH, such as i15:0, 16:1ω7t, 10Me17:0 and cy19:0. In addition, N deposition had an effect on the PLFAs 16:1ω7c and 16:1ω9 (negatively) and cy19:0 (positively). The production of EM mycelia is probably more sensitive to N deposition than total fungal biomass according to the fungal biomarker PLFA 18:2ω6,9. Low amounts of EM mycelia covaried with increased nitrate leaching, suggesting that EM mycelia possibly play an important role in forest soil N retention at increased N input.     

Isotopic signature of nitrate in two contrasting watersheds of Brush Brook,Vermont, USA

We used the dual isotope method to study differences in nitrate export in two subwatersheds in Vermont, USA. Precipitation, soil water and streamwater samples were collected from two watersheds in Camels Hump State Forest, located within the Green Mountains of Vermont. These samples were analyzed for the δ¹⁵N and δ¹⁸O of NO₃⁻. The range of δ¹⁵N–NO₃⁻ values overlapped, with precipitation −4.5‰ to +2.0‰ (n = 14), soil solution −10.3‰ to +6.2‰ (n = 12) and streamwater +0.3‰ to +3.1‰ (n = 69). The δ¹⁸O of precipitation NO₃⁻ (mean 46.8 ± 11.5‰) was significantly different (P < 0.001) from that of the stream (mean 13.2 ± 4.3‰) and soil waters (mean 14.5 ± 4.2‰) even during snowmelt periods. Extracted soil solution and streamwater δ¹⁸O of NO₃⁻ were similar and within the established range of microbially produced NO₃⁻, demonstrating that NO₃⁻ was formed by microbial processes. The δ¹⁵N and δ¹⁸O of NO₃⁻ suggests that although the two tributaries have different seasonal NO₃⁻ concentrations, they have a similar NO₃⁻ source.     

Microbial Cycling of C and N in Northern Hardwood Forests Receiving Chronic Atmospheric NO 3 − Deposition

Sugar maple (Acer saccharum Marsh.)-dominated northern hardwood forests in the upper Lakes States region appear to be particularly sensitive to chronic atmospheric NO₃⁻ deposition. Experimental NO₃⁻ deposition (3 g NO₃⁻ N m⁻² y⁻¹) has significantly reduced soil respiration and increased the export of DOC/DON and NO₃⁻ across the region. Here, we evaluate the possibility that diminished microbial activity in mineral soil was responsible for these ecosystem-level responses to NO₃⁻ deposition. To test this alternative, we measured microbial biomass, respiration, and N transformations in the mineral soil of four northern hardwood stands that have received 9 years of experimental NO₃⁻ deposition. Microbial biomass, microbial respiration, and daily rates of gross and net N transformations were not changed by NO₃⁻ deposition. We also observed no effect of NO₃⁻ deposition on annual rates of net N mineralization. However, NO₃⁻ deposition significantly increased (27%) annual net nitrification, a response that resulted from rapid microbial NO₃⁻ assimilation, the subsequent turnover of NH₄⁺, and increased substrate availability for this process. Nonetheless, greater rates of net nitrification were insufficient to produce the 10-fold observed increase in NO₃⁻ export, suggesting that much of the exported NO₃⁻ resulted directly from the NO₃⁻ deposition treatment. Results suggest that declines in soil respiration and increases in DOC/DON export cannot be attributed to NO₃⁻-induced physiological changes in mineral soil microbial activity. Given the lack of response we have observed in mineral soil, our results point to the potential importance of microbial communities in forest floor, including both saprotrophs and mycorrhizae, in mediating ecosystem-level responses to chronic NO₃⁻ deposition in Lake States northern hardwood forests.     

Effect of C3–C4 Vegetation Change on δ13C and δ15N Values of Soil Organic Matter Fractions Separated by Thermal Stability

Carbon isotopic composition of soils subjected to C₃–C₄ vegetation change can be used to estimate C turnover in bulk soil and in soil organic matter (SOM) pools with fast and intermediate turnover rates. We hypothesized that the biological availability of SOM pools is inversely proportional to their thermal stability, so that thermogravimetry can be used to separate SOM pools with contrasting turnover rates. Soil samples from a field plot cultivated for 10.5 years with the perennial C₄ plant Miscanthus×gigantheus were analyzed by thermogravimetry coupled with differential scanning calorimetry (DSC). Three SOM fractions were distinguished according to the differential weight losses and exothermic or endothermic reactions measured by DSC. The δ¹³C and δ¹⁵N values of these three fractions obtained by gradual soil heating were measured by IRMS. The weight losses up to 190 °C mainly reflected water evaporation because no significant C and N losses were detected and δ¹³C and δ¹⁵N values of the residual SOM remained unchanged. The δ¹³C values (−16.4‰) of SOM fraction decomposed between 190 and 390 °C (containing 79% of total soil C) were slightly closer to that of the Miscanthus plant tissues (δ¹³C = −11.8‰) compared to the δ¹³C values (−16.8‰) of SOM fraction decomposed above 390 °C containing the residual 21% of SOM. Thus, the C turnover in the thermally labile fraction was faster than that in thermally stable fractions, but the differences were not very strong. Therefore, in this first study combining TG-DSC with isotopic analysis, we conclude that the thermal stability of SOM was not very strongly related to biological availability of SOM fractions. In contrast to δ¹³C, the δ¹⁵N values strongly differed between SOM fractions, suggesting that N turnover in the soil was different from C turnover. More detailed fractionation of SOM by thermal analysis with subsequent isotopic analysis may improve the resolution for δ¹³C.     

δ<Superscript>15</Superscript>N constraints on long-term nitrogen balances in temperate forests

Biogeochemical theory emphasizes nitrogen (N) limitation and the many factors that can restrict N accumulation in temperate forests, yet lacks a working model of conditions that can promote naturally high N accumulation. We used a dynamic simulation model of ecosystem N and δ¹⁵N to evaluate which combination of N input and loss pathways could produce a range of high ecosystem N contents characteristic of forests in the Oregon Coast Range. Total ecosystem N at nine study sites ranged from 8,788 to 22,667 kg ha⁻¹ and carbon (C) ranged from 188 to 460 Mg ha⁻¹, with highest values near the coast. Ecosystem δ¹⁵N displayed a curvilinear relationship with ecosystem N content, and largely reflected mineral soil, which accounted for 96–98% of total ecosystem N. Model simulations of ecosystem N balances parameterized with field rates of N leaching required long-term average N inputs that exceed atmospheric deposition and asymbiotic and epiphytic N₂-fixation, and that were consistent with cycles of post-fire N₂-fixation by early-successional red alder. Soil water δ¹⁵NO₃ ⁻ patterns suggested a shift in relative N losses from denitrification to nitrate leaching as N accumulated, and simulations identified nitrate leaching as the primary N loss pathway that constrains maximum N accumulation. Whereas current theory emphasizes constraints on biological N₂-fixation and disturbance-mediated N losses as factors that limit N accumulation in temperate forests, our results suggest that wildfire can foster substantial long-term N accumulation in ecosystems that are colonized by symbiotic N₂-fixing vegetation.     

The effect of forest type on throughfall deposition and seepage flux: a review

Converting deciduous forests to coniferous plantations and vice versa causes environmental changes, but till now insight into the overall effect is lacking. This review, based on 38 case studies, aims to find out how coniferous and deciduous forests differ in terms of throughfall (+stemflow) deposition and seepage flux to groundwater. From the comparison of coniferous and deciduous stands at comparable sites, it can be inferred that deciduous forests receive less N and S via throughfall (+stemflow) deposition on the forest floor. In regions with relatively low open field deposition of atmospheric N (<10 kg N ha⁻¹ year⁻¹), lower NH₄⁺ mean throughfall (+stemflow) deposition was, however, reported under conifers compared to deciduous forest, while in regions with high atmospheric N pollution (>10 kg N ha⁻¹ year⁻¹), the opposite could be concluded. The higher the open field deposition of NH₄⁺, the bigger the difference between the coniferous and deciduous throughfall (+stemflow) deposition. Furthermore, it can be concluded that canopy exchange of K⁺, Ca²⁺ and Mg²⁺ is on average higher in deciduous stands. The significantly higher stand deposition flux of N and S in coniferous forests is reflected in a higher soil seepage flux of NO₃⁻, SO₄²⁻, K⁺, Ca²⁺, Mg²⁺ and Al(III). Considering a subset of papers for which all necessary data were available, a close relationship between throughfall (+stemflow) deposition and seepage was found for N, irrespective of the forest type, while this was not the case for S. This review shows that the higher input flux of N and S in coniferous forests clearly involves a higher seepage of NO₃⁻ and SO₄²⁻ and accompanying cations K⁺, Ca²⁺, Mg²⁺ and Al(III) into the groundwater, making this forest type more vulnerable to acidification and eutrophication compared to the deciduous forest type.     

Abiotic immobilization of nitrate in two soils of relic <Emphasis Type="Italic">Abies pinsapo</Emphasis>-fir forests under Mediterranean climate

Evidence for abiotic immobilization of nitrogen (N) in soil is accumulating, but remains controversial. Identifying the fate of N from atmospheric deposition is important for understanding the N cycle of forest ecosystems. We studied soils of two Abies pinsapo fir forests under Mediterranean climate seasonality in southern Spain—one with low N availability and the other with symptoms of N saturation. We hypothesized that biotic and abiotic immobilization of nitrate (NO₃ ⁻) would be lower in soils under these forests compared to more mesic temperate forests, and that the N saturated stand would have the lowest rates of NO₃ ⁻ immobilization. Live and autoclaved soils were incubated with added ¹⁵NO₃ ⁻ (10 μg N g⁻¹ dry soil; 99% enriched) for 24 h, and the label was recovered as total dissolved-N, NO₃ ⁻, ammonium (NH₄ ⁺), or dissolved organic-N (DON). To evaluate concerns about possible iron interference in analysis of NO₃ ⁻ concentrations, both flow injection analysis (FIA) and ion chromatography (IC) were applied to water extracts, soluble iron was measured in both water and salt extracts, and standard additions of NO₃ ⁻ to salt extracts were analyzed. Good agreement between FIA and IC analysis, low concentrations of soluble Fe, and 100% (±3%) recovery of NO₃ ⁻ standard additions all pointed to absence of an interference problem for NO₃ ⁻ quantification. On average, 85% of the added ¹⁵NO₃ ⁻ label was recovered as ¹⁵NO₃ ⁻, which supports our hypothesis that rates of immobilization were generally low in these soils. A small amount (mean = 0.06 μg N g⁻¹ dry soil) was recovered as ¹⁵NH₄ ⁺ in live soils and none in sterilized soils. Mean recovery as DO¹⁵N ranged from 0.6 to 1.5 μg N g⁻¹ dry soil, with no statistically significant effect of sterilization or soil type, indicating that this was an abiotic process that occurred at similar rates in both soils. These results demonstrate a detectable, but modest rate of abiotic immobilization of NO₃ ⁻ to DON, supporting our first hypothesis. These mineral soils may not have adequate carbon availability to support the regeneration of reducing microsites needed for high rates of NO₃ ⁻ reduction. Our second hypothesis regarding lower expected abiotic immobilization in soils from the N-saturated site was not supported. The rates of N deposition in this region may not be high enough to have swamped the capacity for soil NO₃ ⁻ immobilization, even in the stand showing some symptoms of N saturation. A growing body of evidence suggests that soil abiotic NO₃ ⁻ immobilization is common, but that rates are influenced by a combination of factors, including the presence of plentiful available carbon, reduced minerals in anaerobic microsites and adequate NO₃ ⁻ supply.     

Plant and Soil N Response of Southern Californian Semi-arid Shrublands After 1 Year of Experimental N Deposition

Large inputs of atmospheric N from dry deposition accumulate on vegetation and soil surfaces of southern Californian chaparral and coastal sage scrub (CSS) ecosystems during the late-summer and early-fall and become available as a pulse following winter rainfall; however, the fate of this dry season atmospheric N addition is unknown. To assess the potential for dry season atmospheric N inputs to be incorporated into soil and/or vegetation N pools, an in situ N addition experiment was initiated in a post-fire chaparral and a mature CSS stand where 10 × 10 m plots were exposed to either ambient N deposition (control) or ambient +50 kg N ha⁻¹ (added N) added as NH₄NO₃ during a single application in October 2003. After 1 year of N addition, plots exposed to added N had significantly higher accumulation of extractable inorganic N (NH₄−N + NO₃−N) on ion exchange resins deployed in the 0–10 cm mineral soil layer and higher soil extractable N in the subsurface (30–40 cm) mineral soil than plots exposed to ambient N. Chaparral and CSS shrubs exposed to added N also exhibited a significant increase in tissue N concentration and a decline in the tissue C:N ratio, and added N significantly altered the shrub tissue δ ¹⁵N natural abundance. Leaching of inorganic N to 1 m below the soil surface was on average 2–3 times higher in the added N plots, but large within treatment variability cause these differences to be statistically insignificant. Although a large fraction of the added N could not be accounted for in the shrub and soil N pools investigated, these observations suggest that dry season N inputs can significantly and rapidly alter N availability and shrub tissue chemistry in Mediterranean-type chaparral and CSS shrublands of southern California.     

The chemical composition of throughfall beneath oak, birch and pine canopies in Northwest Germany

The chemical composition of rainwater is altered upon its passage through tree canopies. In order to investigate how rainwater chemistry is affected by canopy-dependent processes in characteristic forest types of Northwest German sandy lowland regions – oak–birch-forests, Betula pubescens Ehrh. swamp forests, and stands of Pinus sylvestris L. – comparative studies on the chemical composition of throughfall were carried out at seven forest sites, situated in close proximity within a nature reserve. Additionally, rainwater was sampled at three heathland sites for analysis of open-field precipitation and at three sites along an oak–birch-forest edge. Throughfall concentrations of most of the parameters analysed were significantly higher than open-field concentrations, especially with regard to electric conductivity, NH₄-N, K⁺, and KMnO₄-index. Ion concentrations in throughfall were the lowest in a 10-year-old stand of Betula pendula Roth. and Pinus sylvestris and in a Betula pubescens swamp forest and were highest beneath a stand of Pinus sylvestris. Except for Na⁺, Cl⁻, and NO₃⁻, ion concentrations in both throughfall and open-field precipitation increased during the growing season (May–October). In throughfall, Ca²⁺, Mg²⁺, K⁺, and Mn²⁺ were strongly correlated. Enrichment ratios between throughfall and open-field deposition varied among sites and elements and were the highest for K^‰+, Mg^2‰+, and Mn^2‰+. Estimates of canopy leaching indicated high leaching rates of K^‰+ and Mn^2‰+ and moderate leaching of Mg^2‰+. The contribution of foliar leaching to throughfall deposition was higher at the deciduous than at the coniferous stands.     

Estimation of symbiotically fixed nitrogen in field grown soybeans: An application of natural15N/14N abundance and a low level15N-tracer technique

Summary Plants from agricultural and natural upland ecosystem were investigated for¹⁵N content to evaluate the role of symbiotic N₂-fixation in the nitrogen nutrition of soybean. Increased yields and lower δ¹⁵N values of nodulating soybeansvs, non-nodulating isolines gave semi-quantitative estimates of N₂ fixation. A fairly large discrepancy was found between estimations by δ¹⁵N and by N yield at 0 kg N/ha of fertilizer. More precise estimates were made by following changes in plant δ¹⁵N when fertilizer δ¹⁵N was varied near¹⁵N natural abundance level. Clearcut linear relationships between δ¹⁵N values of whole plants and of fertilizer were obtained at 30 kg N/ha of fertilizer for three kinds of soils. In experimental field plots, nodulating soybeans obtained 13±1% of their nitrogen from fertilizer, 66±8% from N₂ fixation and 21±10% from soil nitrogen in Andosol brown soil; 30%, 16% and 54% in Andosol black soil; 7%, 77% and 16% in Alluvial soil, respectively. These values for N₂ fixation coincided with each corresponding estimation by N yield method. Other results include: 1)¹⁵N content in upland soils and plants was variable, and may reflect differences in the mode of mineralization of soil organics, and 2) nitrogen isotopic discrimination during fertilizer uptake (δ¹⁵N of plant minus fertilizer) ranged from −2.2 to +4.9‰ at 0–30 kg N/ha of fertilizer, depending on soil type and plant species. The proposed method can accurately and relatively simply establish the importance of symbiotic nitrogen fixation for soybeans growing in agricultural settings.     

Nitrogen availability patterns in white-sand vegetations of Central Brazilian Amazon

Addressing spatial variability in nitrogen (N) availability in the Central Brazilian Amazon, we hypothesized that N availability varies among white-sand vegetation types (campina and campinarana) and lowland tropical forests (dense terra-firme forests) in the Central Brazilian Amazon, under the same climate conditions. Accordingly, we measured soil and foliar N concentration and N isotope ratios (δ¹⁵N) throughout the campina-campinarana transect and compared to published dense terra-firme forest results. There were no differences between white-sand vegetation types in regard to soil N concentration, C:N ratio and δ¹⁵N across the transect. Both white-sand vegetation types showed very low foliar N concentrations and elevated foliar C:N ratios, and no significant difference between site types was observed. Foliar δ¹⁵N was depleted, varying from −9.6 to 1.6‰ in the white-sand vegetations. The legume Aldina heterophylla had the highest average δ¹⁵N values (−1.5‰) as well as the highest foliar N concentration (2.1%) while the non-legume species had more depleted δ¹⁵N values and the average foliar N concentrations varied from 0.9 to 1.5% among them. Despite the high variation in foliar δ¹⁵N among plants, a significant and gradual ¹⁵N-enrichment in foliar isotopic signatures throughout the campina–campinarana transect was observed. Individual plants growing in the campinarana were significantly enriched in ¹⁵N compared to those in campina. In the white-sand N-limited ecosystems, the differentiation of N use seems to be a major cause of variations observed in foliar δ¹⁵N values throughout the campina–campinarana transect.     

Large Loss of Dissolved Organic Nitrogen from Nitrogen-Saturated Forests in Subtropical China

Dissolved organic nitrogen (DON) has recently been recognized as an important component of terrestrial N cycling, especially under N-limited conditions; however, the effect of increased atmospheric N deposition on DON production and loss from forest soils remains controversial. Here we report DON and dissolved organic carbon (DOC) losses from forest soils receiving very high long-term ambient atmospheric N deposition with or without additional experimental N inputs, to investigate DON biogeochemistry under N-saturated conditions. We studied an old-growth forest, a young pine forest, and a young mixed pine/broadleaf forest in subtropical southern China. All three forests have previously been shown to have high nitrate (NO₃⁻) leaching losses, with the highest loss found in the old-growth forest. We hypothesized that DON leaching loss would be forest specific and that the strongest response to experimental N input would be in the N-saturated old-growth forest. Our results showed that under ambient deposition (35–50 kg N ha⁻¹ y⁻¹ as throughfall input), DON leaching below the major rooting zone in all three forests was high (6.5–16.9 kg N ha⁻¹ y⁻¹). DON leaching increased 35–162% following 2.5 years of experimental input of 50–150 kg N ha⁻¹ y⁻¹. The fertilizer-driven increase of DON leaching comprised 4–17% of the added N. A concurrent increase in DOC loss was observed only in the pine forest, even though DOC:DON ratios declined in all three forests. Our data showed that DON accounted for 23–38% of total dissolved N in leaching, highlighting that DON could be a significant pathway of N loss from forests moving toward N saturation. The most pronounced N treatment effect on DON fluxes was not found in the old-growth forest that had the highest DON loss under ambient conditions. DON leaching was highly correlated with NO₃⁻ leaching in all three forests. We hypothesize that abiotic incorporation of excess NO₃⁻ (through chemically reactive NO₂⁻) into soil organic matter and the consequent production of N-enriched dissolved organic matter is a major mechanism for the consistent and large DON loss in the N-saturated subtropical forests of southern China. Dr. YT Fang performed research, analyzed data, and wrote the paper; Prof. WX Zhu participated in the initial experimental design, analyzed data, and took part in writing the paper; Prof. P Gundersen conceived the study and took part in writing; Prof. JM Mo and Prof. GY Zhou conceived study; Prof. M Yoh analyzed part of the data and contributed to the development of DON model.     

Tree Patches Show Greater N Losses but Maintain Higher Soil N Availability than Grassland Patches in a Frequently Burned Oak Savanna

Long-term prescribed fires have increased woody canopy openness and reduced nitrogen (N) cycling (that is, net N mineralization) in an oak savanna in Minnesota, USA. It is unclear how fire-induced shifts from oak-dominated to C4 grass-dominated vegetation contribute to this decline in N cycling compared to direct effects of increasing fire frequency promoting greater N losses. We determined (1) the magnitude of decline in net N mineralization in oak versus grass-dominated patches with increasing fire frequency and (2) if differences in net N mineralization between oak and grass patches in frequently burned oak savanna (burned 8 out of 10 years on average during the last 40 years) could be attributed to differences in N losses through volatilization and leaching or to plant traits affecting decomposition and mineralization. In situ net N mineralization declined with increasing fire frequency overall, but this decline was less in oak- than in grass-dominated patches, with oak-dominated patches having more than two times higher net N mineralization than grass-dominated patches. Greater net N mineralization in oak-dominated patches occurred despite greater N losses through volatilization and leaching (on average 1.8 and 1.4 g m⁻² y⁻¹ for oak- and grass-dominated patches, respectively), likely because of higher plant litter N concentration in the oak-dominated patches. As total soil N pools in the first 15 cm did not differ between oak- and grass-dominated patches (on average 83 g N m⁻²), N inputs from atmospheric deposition and uptake from deep soil layers may offset higher N losses. Our results further show that net N mineralization rates decline within 5 years after tree death and subsequent colonization by C4 grasses to levels observed in grass-dominated patches. Although long-term prescribed fires often directly reduce N stocks and cycling because of increased N losses, this study has shown that fire-induced shifts in vegetation composition can strongly contribute to the declines in N cycling in systems that are frequently disturbed by fires with potential feedbacks to plant productivity.     

Soil Inorganic N Leaching in Edges of Different Forest Types Subject to High N Deposition Loads

We report on soil leaching of dissolved inorganic nitrogen (DIN) along transects across exposed edges of four coniferous and four deciduous forest stands. In a 64-m edge zone, DIN leaching below the main rooting zone was enhanced relative to the interior (at 128 m from the edge) by 21 and 14 kg N ha⁻¹ y⁻¹ in the coniferous and deciduous forest stands, respectively. However, the patterns of DIN leaching did not univocally reflect those of DIN throughfall deposition. DIN leaching in the first 20 m of the edges was lower than at 32–64 m from the edge (17 vs. 36 kg N ha⁻¹ y⁻¹ and 15 vs. 24 kg N ha⁻¹ y⁻¹ in the coniferous and deciduous forests, respectively). Nitrogen stocks in the mineral topsoil (0–30 cm) were, on average, 943 kg N ha⁻¹ higher at the outer edges than in the interior, indicating that N retention in the soil is probably one of the processes involved in the relatively low DIN leaching in the outer edges. We suggest that a complex of edge effects on biogeochemical processes occurs at the forest edges as a result of the interaction between microclimate, tree dynamics (growth and litterfall), and atmospheric deposition of N and base cations.     

Plant and microbial nitrogen use and turnover: Rapid conversion of nitrate to ammonium in soil with roots

Immobilization of ammonium (NH ₄ ⁺ ) by plants and microbes, a controlling factor of ecosystem nitrogen (N) retention, has usually been measured based on uptake of¹⁵NH ₄ ⁺ solutions injected into soil. To study the influence of roots on N dynamics without stimulating consumption of NH ₄ ⁺ , we estimated gross nitrification in the presence or absence of live roots in an agricultural soil. Tomato (Lycopersicon esculentum var. Peto76) plants were grown in microcosms containing root exclosures. When the plants were 7 weeks old,¹⁵N enriched nitrate (NO ₃ ⁻ ) was applied in the 0–150 mm soil layer. After 24 h, > 30 times more¹⁵NH ₄ ⁺ was found in the soil with roots than in the soil of the root exclosures. At least 18% of the NH ₄ ⁺ -N present at this time in the soil with roots had been converted from NO ₃ ⁻ . We estimated rates of conversion of NO ₃ ⁻ to NH ₄ ⁺ , and rates ofNH ₄ ⁺ immobilization by plants and microbes, by simulating N-flow of¹⁴⁺¹⁵N and¹⁵N in three models representing mechanisms that may be underlying the experimental data: Dissimilatory NO ₃ ⁻ reduction to NH ₄ ⁺ (DNRA), plant N efflux, and microbial biomass nitrogen (MBN) turnover. Compared to NO ₃ ⁻ uptake, plant NH ₄ ⁺ uptake was modest. Ammonium immobilization by plants and microbes was equal to at least 35% of nitrification rates. The rapid recycling of NO ₃ ⁻ to NH ₄ ⁺ via plants and/or microbes contributes to ecosystem N retention and may enable plants growing in agricultural soils to capture more NH ₄ ⁺ than generally assumed.