Nitrogen Distribution and Cycling through Water Flows in a Subtropical Bamboo Forest under High Level of Atmospheric Deposition

Background

The hydrological cycle is an important way of transportation and reallocation of reactive nitrogen (N) in forest ecosystems. However, under a high level of atmospheric N deposition, the N distribution and cycling through water flows in forest ecosystems especially in bamboo ecosystems are not well understood.

Methodology/Principal Findings

In order to investigate N fluxes through water flows in a Pleioblastus amarus bamboo forest, event rainfall/snowfall (precipitation, PP), throughfall (TF), stemflow (SF), surface runoff (SR), forest floor leachate (FFL), soil water at the depth of 40 cm (SW1) and 100 cm (SW2) were collected and measured through the whole year of 2009. Nitrogen distribution in different pools in this ecosystem was also measured. Mean N pools in vegetation and soil (0–1 m) were 351.7 and 7752.8 kg ha⁻¹. Open field nitrogen deposition at the study site was 113.8 kg N ha⁻¹ yr⁻¹, which was one of the highest in the world. N-NH₄ ⁺, N-NO₃ ⁻ and dissolved organic N (DON) accounted for 54%, 22% and 24% of total wet N deposition. Net canopy accumulated of N occurred with N-NO₃ ⁻ and DON but not N-NH₄ ⁺. The flux of total dissolved N (TDN) to the forest floor was greater than that in open field precipitation by 17.7 kg N ha⁻¹ yr⁻¹, due to capture of dry and cloudwater deposition net of canopy uptake. There were significant negative exponential relationships between monthly water flow depths and monthly mean TDN concentrations in PP, TF, SR, FFL and SW1.

Conclusions/Significance

The open field nitrogen deposition through precipitation is very high over the world, which is the main way of reactive N input in this bamboo ecosystem. The water exchange and N consume mainly occurred in the litter floor layer and topsoil layer, where most of fine roots of bamboo distributed.     

Spatial pattern of soil nitrogen availability and its relationship to stand structure in a coniferous-broadleaved mixed forest with a dense dwarf bamboo understory in northern Japan

Natural disturbances create spatial patterns of the ecosystem processes and functions in natural forests. However, how dynamics and the spatial structure of forests relate to soil nitrogen dynamics is not well understood. We examined the spatial relationship between the distributions of canopy and understory species, and soil nitrogen dynamics in a natural coniferous-broadleaved mixed forest with a dense understory of Sasa dwarf bamboo in northern Japan. The O horizon was thick where coniferous litter predominated, and it was thin where broadleaved litter predominated. The soil water content was low in areas with a thick O horizon and a high abundance of coniferous trees. The soil nitrate content was low where the soil water content was low, and the soil nitrate content increased linearly with increasing net nitrification potential. These results suggest that the soil nitrate content under the coniferous canopy was lower because of the low nitrification potential of soil microbes in soils with low water contents. The soil nitrate content and nitrification potential were higher in the canopy gap than under the canopy. Our results suggest that forest structure, specifically the thickness of the forest floor, significantly affects the spatial pattern of the soil water content, thereby creating a spatial pattern of soil nitrogen availability at a relatively small scale with flat topography. The higher nitrification potential under the canopy gap could pose a long-term risk of nitrate leaching because of the suppression of the natural regeneration of canopy species by dense Sasa dwarf bamboo in this forest ecosystem.     

Soil nitrogen turnover in proximal and distal stem areas of European beech trees

In European beech (Fagus sylvatica L.) forests, a large proportion of the water and ion input to the soil results from stemflow which creates a soil microsite of high element fluxes proximal to the tree trunk. The soil proximal to the stem is considered to have different rates of nitrogen turnover which might influence the estimation of N-turnover rates at the stand scale. In a previous study we reported high nitrate fluxes with seepage proximal to the stems in a forest dominated by European beech in Steigerwald, Germany. Here, we investigated the soil nitrogen turnover in the top 15 cm soil in proximal (defined as 1 m² around beech stems) and distal stem areas. Laboratory incubations and in situ sequential coring incubations were used to determine the net rates of ammonification, nitrification, and root uptake of mineral nitrogen. In the laboratory incubations higher rates of net nitrogen mineralization and nitrification were found in the forest floor proximal to the stem as compared to distal stem areas. No stem related differences were observed in case of mineral soil samples. In contrast, the in situ incubations revealed higher rates of nitrification in the mineral soil in proximal stem areas, while net nitrogen mineralization was equal in proximal and distal areas. In the in situ incubations the average ratio of nitrification/ammonification was 0.85 in proximal and 0.34 in distal stem areas. The net nitrogen mineralization was 4.4 g N m^-2 90 day^-1 in both areas. Mineralized nitrogen was almost completely taken up by tree roots with ammonium as the dominant nitrogen species. The average ratio of nitrate/ammonium uptake was 0.69 in proximal and 0.20 in distal areas. The higher water content of the soil in proximal stem areas is considered to be the major reason for the increased rates of nitrification. Different nitrogen turnover rates in proximal stem areas had no influence on the nitrogen turnover rates in soil at the stand scale. Consequently, the observed high nitrate fluxes with seepage proximal to stems are attributed to the high nitrogen input by stemflow rather than to soil nitrogen turnover. This revised version was published online in August 2006 with corrections to the Cover Date.     

Effects of plant diversity,N fertilization,and elevated carbon dioxide on grassland soil N cycling in a long‐term experiment

The effects of global environmental changes on soil nitrogen (N) pools and fluxes have consequences for ecosystem functions such as plant productivity and N retention. In a 13‐year grassland experiment, we evaluated how elevated atmospheric carbon dioxide (CO₂), N fertilization, and plant species richness alter soil N cycling. We focused on soil inorganic N pools, including ammonium and nitrate, and two N fluxes, net N mineralization and net nitrification. In contrast with existing hypotheses, such as progressive N limitation, and with observations from other, often shorter, studies, elevated CO₂ had relatively static and small, or insignificant, effects on soil inorganic N pools and fluxes. Nitrogen fertilization had inconsistent effects on soil N transformations, but increased soil nitrate and ammonium concentrations. Plant species richness had increasingly positive effects on soil N transformations over time, likely because in diverse subplots the concentrations of N in roots increased over time. Species richness also had increasingly positive effects on concentrations of ammonium in soil, perhaps because more carbon accumulated in soils of diverse subplots, providing exchange sites for ammonium. By contrast, subplots planted with 16 species had lower soil nitrate concentrations than less diverse subplots, especially when fertilized, probably due to greater N uptake capacity of subplots with 16 species. Monocultures of different plant functional types had distinct effects on N transformations and nitrate concentrations, such that not all monocultures differed from diverse subplots in the same manner. The first few years of data would not have adequately forecast the effects of N fertilization and diversity on soil N cycling in later years; therefore, the dearth of long‐term manipulations of plant species richness and N inputs is a hindrance to forecasting the state of the soil N cycle and ecosystem functions in extant plant communities.     

Is there synchronicity in nitrogen input and output fluxes at the Noland Divide Watershed,a small N-saturated forested catchment in the Great Smoky Mountains National Park?

High-elevation red spruce [Picea rubens Sarg.]-Fraser fir [Abies fraseri (Pursh.) Poir] forests in the Southern Appalachians currently receive large nitrogen (N) inputs via atmospheric deposition (30 kg N ha(-1) year(-1)) but have limited N retention capacity due to a combination of stand age, heavy fir mortality caused by exotic insect infestations, and numerous gaps caused by windfalls and ice storms. This study examined the magnitude and timing of the N fluxes into, through, and out of a small, first-order catchment in the Great Smoky Mountains National Park. It also examined the role of climatic conditions in causing interannual variations in the N output signal. About half of the atmospheric N input was exported annually in the streamwater, primarily as nitrate (NO3-N). While most incoming ammonium (NH4-N) was retained in the canopy and the forest floor, the NO3-N fluxes were very dynamic in space as well as in time. There was a clear decoupling between NO3-N input and output fluxes. Atmospheric N input was greatest in the growing season while largest NO3-N losses typically occurred in the dormant season. Also, as water passed through the various catchment compartments, the NO3-N flux declined below the canopy, increased in the upper soil due to internal N mineralization and nitrification, and declined again deeper in the mineral soil due to plant uptake and microbial processing. Temperature control on N production and hydrologic control on NO3-N leaching during the growing season likely caused the observed inter-annual variation in fall peak NO3-N concentrations and N discharge rates in the stream.     

Insect herbivory, organic matter deposition and effects on belowground organic matter fluxes in a central European oak forest

Apart from the forest floor, the canopy of forested ecosystems functions as the second most important source for dissolved and particulate fractions of organic and inorganic C and N compounds. However, under mass outbreak situations of insect herbivores this flux path of organic matter is considerably intensified clearly exceeding C and N fluxes from the forest floor. In this paper we report on herbivore-altered C and N fluxes from the canopy to the forest floor and effects on forest floor nutrient fluxes during severe defoliating herbivory of the winter moth (Operophtera brumata) and the mottled umber moth (Eranis defoliaria) in an oak forest in Germany. Over the course of 6.5 months we followed the C and N fluxes with bulk deposition, throughfall solution, insect frass deposits (green-fall together with insect faeces) and with forest floor solution in an 117-yr-old oak (Quercus petraea) forest. Compared to the control, herbivore defoliation significantly enhanced throughfall inputs of total and dissolved organic carbon and nitrogen by a factor of 3 and 2.5 (for TOC and DOC), and by 1.4 and 1.3 times (for TNb and DNb), respectively. Frass plus green-fall C and N fluxes peaked in May with 592 kg C?ha^?1 and 33.5 kg N?ha^?1 representing 79.6% (for C) and 78.3% (for N) of the total C and N input over 2.5 months. The quantitative and qualitative C and N input via faeces and litter deposition significantly differ between the insect affected and non-affected site. However, the C and N fluxes with throughfall did not significantly correlate with forest floor leachates. In this context, forest floor fluxes of TOC, DOC and NO₃-N were significantly lower at the infested site compared to the control, whereas fluxes of NH₄-N together with DON were significantly higher. The study demonstrates the importance of linking the population and associated frass dynamics of herbivorous insects with the cycling of nutrients and organic matter in forest ecosystems, highlighting the remarkable alterations in the timing, amounts and nature of organic matter dynamics on the ecosystem level. Consequently, the ecology of phytophagous insects allows partly to explain temporal-spatial alterations in nutrient cycling and thus ecosystem functioning.     

Rainfall input, throughfall and stemflow of nutrients in a central African rain forest dominated by ectomycorrhizal trees

Incident rainfall is a major source of nutrient input to a forest ecosystem and the consequent throughfall and stemflow contribute to nutrient cycling. These rain-based fluxes were measured over 12 mo in two forest types in Korup National Park, Cameroon, one with low (LEM) and one with high (HEM) ectomycorrhizal abundances of trees. Throughfall was 96.6 and 92.4% of the incident annual rainfall (5370 mm) in LEM and HEM forests respectively; stemflow was correspondingly 1.5 and 2.2%. Architectural analysis showed that ln(funneling ratio) declined linearly with increasing ln(basal area) of trees. Mean annual inputs of N, P, K, Mg and Ca in incident rainfall were 1.50, 1.07, 7.77, 5.25 and 9.27 kg ha^–1, and total rain-based inputs to the forest floor were 5.0, 3.2, 123.4, 14.4 and 37.7 kg ha^–1 respectively. The value for K is high for tropical forests and that for N is low. Nitrogen showed a significantly lower loading of throughfall and stemflow in HEM than in LEM forest, this being associated in the HEM forest with a greater abundance of epiphytic bryophytes which may absorb more N. Incident rainfall provided c. 35% of the gross input of P to the forest floor (i.e., rain-based plus small litter inputs), a surprisingly high contribution given the sandy P-poor soils. At the start of the wet season leaching of K from the canopy was particularly high. Calcium in the rain was also highest at this time, most likely due to washing off of dry-deposited Harmattan dusts. It is proposed that throughfall has an important 'priming' function in the rapid decomposition of litter and mineralization of P at the start of the wet season. The contribution of P inputted from the atmosphere appears to be significant when compared to the rates of P mineralization from leaf litter.     

Growth,Net Production,Litter Decomposition,and Net Nitrogen Accumulation by Epiphytic Bryophytes in a Tropical Montane Forest1

To understand the ecological roles of epiphytic bryophytes in the carbon (C) and nitrogen (N) cycles of a tropical montane forest, we used samples in enclosures to estimate rates of growth, net production, and N accumulation by shoots in the canopy, and litterbags, to estimate rates of decomposition and N dynamics of epiphytic bryophyte litter in the canopy and on the forest floor in Monteverde, Costa Rica. Growth of epiphytic bryophytes was estimated at 30.0–49.9 percent/yr, net production at 122–203 g/m²/yr, and N accumulation at 1.8–3.0 g N/m²/yr. Cumulative mass loss from litterbags after one and two years in the canopy was 17 ± 2 and 19 ± 2 percent (mean ± 1 SE) of initial sample mass, respectively, and mass loss from litter and green shoots in litterbags after one year on the forest floor was 29 ± 2 and 45 ± 3 percent, respectively. Approximately 30 percent of the initial N mass was released rapidly from litter in both locations. Nitrogen loss from green shoots on the forest floor was greater; about 47 percent of the initial N mass was lost within the first three months. There was no evidence for net N immobilization by litter or green shoots, but the remaining N in litter was apparently recalcitrant. Annual net accumulation of C and N by epiphytic bryophytes was estimated at 37–64 g C/m²/yr and 0.8–1.3 g N/m²/yr, respectively. Previous research at this site indicated that epiphytic bryophytes retain inorganic N from atmospheric deposition to the canopy. Therefore, they play a major role in transforming N from mobile to highly recalcitrant forms in this ecosystem.     

The influence of western hemlock and western redcedar on microbial numbers,nitrogen mineralization,and nitrification

Summary Microbial numbers in the forest floor and mineral soil (Al horizon) under large individual western hemlock (Tsuga heterophylla) and western redcedar (Thuja plicata) trees were compared. The lower pH and base saturation of hemlock samples was associated with higher fungal spore counts while cedar samples had higher total microbial counts and populations of ammonium oxidizing bacteria. Nitrogen mineralization rates were greater in laboratory incubations of hemlock soil but nitrification was only observed in incubations of cedar soil. These differences in nitrogen mineralization and nitrification are aspects of species-specific nutrient cycling regimes.     

Nitrogen Cycling Responses to Mountain Pine Beetle Disturbance in a High Elevation Whitebark Pine Ecosystem

Ecological disturbances can significantly affect biogeochemical cycles in terrestrial ecosystems, but the biogeochemical consequences of the extensive mountain pine beetle outbreak in high elevation whitebark pine (WbP) (Pinus albicaulis) ecosystems of western North America have not been previously investigated. Mountain pine beetle attack has driven widespread WbP mortality, which could drive shifts in both the pools and fluxes of nitrogen (N) within these ecosystems. Because N availability can limit forest regrowth, understanding how beetle-induced mortality affects N cycling in WbP stands may be critical to understanding the trajectory of ecosystem recovery. Thus, we measured above- and belowground N pools and fluxes for trees representing three different times since beetle attack, including unattacked trees. Litterfall N inputs were more than ten times higher under recently attacked trees compared to unattacked trees. Soil inorganic N concentrations also increased following beetle attack, potentially driven by a more than two-fold increase in ammonium (NH₄ ⁺) concentrations in the surface soil organic horizon. However, there were no significant differences in mineral soil inorganic N or soil microbial biomass N concentrations between attacked and unattacked trees, implying that short-term changes in N cycling in response to the initial stages of WbP attack were restricted to the organic horizon. Our results suggest that while mountain pine beetle attack drives a pulse of N from the canopy to the forest floor, changes in litterfall quality and quantity do not have profound effects on soil biogeochemical cycling, at least in the short-term. However, continuous observation of these important ecosystems will be crucial to determining the long-term biogeochemical effects of mountain pine beetle outbreaks.     

The complete nitrogen cycle of an N-saturated spruce forest ecosystem

Long-term nitrogen deposition into forest ecosystems has turned many forests in Central Europe and North America from N-limited to N-saturated systems, with consequences for climate as well as air and groundwater quality. However, complete quantification of processes that convert the N deposited and contributed to ecosystem N cycling is scarce. In this study, we provide the first complete quantification of external and internal N fluxes in an old-growth spruce forest, the Höglwald, Bavaria, Germany, exposed to high chronic N deposition. In this forest, N cycling is dominated by high rates of mineralisation of soil organic matter, nitrification and immobilisation of ammonium and nitrate into microbial biomass. The amount of ammonium available is sufficient to cover the entire N demand of the spruce trees. The data demonstrate the existence of a highly dynamic internal N cycle within the soil, driven by growth and death of the microbial biomass, which turns over approximately seven times each year. Although input and output fluxes are of high environmental significance, they are low compared to the internal fluxes mediated by microbial activity.     

Response of dissolved organic matter in the forest floor to long-term manipulation of litter and throughfall inputs

Dissolved organic matter (DOM) contributes to organic carbon either stored in mineral soil horizons or exported to the hydrosphere. However, the main controls of DOM dynamics are still under debate. We studied fresh leaf litter and more decomposed organic material as the main sources of DOM exported from the forest floor of a mixed beech/oak forest in Germany. In the field we doubled and excluded aboveground litter input and doubled the input of throughfall. From 1999 to 2005 we measured concentrations and fluxes of dissolved organic C and N (DOC, DON) beneath the Oi and Oe/Oa horizon. DOM composition was traced by UV and fluorescence spectroscopy. In selected DOM samples we analyzed the concentrations of phenols, pentoses and hexoses, and lignin-derived phenols by CuO oxidation. DOC and DON concentrations and fluxes almost doubled instantaneously in both horizons of the forest floor by doubling the litter input and DOC concentrations averaged 82 mg C l⁻¹ in the Oe/Oa horizon. Properties of DOM did not suggest a change of the main DOM source towards fresh litter. In turn, increasing ratios of hexoses to pentoses and a larger content of lignin-derived phenols in the Oe/Oa horizon of the Double litter plots in comparison to the Control plots indicated a priming effect: Addition of fresh litter stimulated microbial activity resulting in increased microbial production of DOM from organic material already stored in Oe/Oa horizons. Exclusion of litter input resulted in an immediate decrease in DOC concentrations and fluxes in the thin Oi horizon. In the Oe/Oa horizon DOC concentrations started to decline in the third year and were significantly smaller than those in the Control after 5 years. Properties of DOM indicated an increased proportion of microbially and throughfall derived compounds after exclusion of litter inputs. Dissolved organic N did not decrease upon litter exclusion. We assume a microbial transformation of mineral N from throughfall and N mineralization to DON. Increased amounts of throughfall resulted in almost equivalently increased DOC fluxes in the Oe/Oa horizon. However, long-term additional throughfall inputs resulted in significantly declining DOC concentrations over time. We conclude that DOM leaving the forest floor derives mainly from decomposed organic material stored in Oe/Oa horizons. Leaching of organic matter from fresh litter is of less importance. Observed effects of litter manipulations strongly depend on time and the stocks of organic matter in forest floor horizons. Long-term experiments are particularly necessary in soils/horizons with large stocks of organic matter and in studies focusing on effects of declined substrate availability. The expected increased primary production upon climate change with subsequently enhanced litter input may result in an increased production of DOM from organic soil horizons.     

A field study on the fate of 15N-ammonium to demonstrate nitrification of atmospheric ammonium in an acid forest soil

To demonstrate the contribution of atmospheric ammonium to soil acidification in acid forest soils, a field study with¹³N-ammonium as tracer was performed in an oak-birch forest soil. Monitoring and analysis of soil solutions from various depths on the¹³N-ammonium and¹⁵N-nitrate contents, showed that about 54% of the applied¹⁵N-ammonium was oxidized to nitrate in the forest floor. Over a period of one year about 20% of the¹⁵N remained as organic nitrogen in this layer. The percentage¹⁵N enrichment in ammonium and nitrate were in the same range in all the forest floor percolates, indicating that even in extremely acid forest soils (pH < 4) nitrate formation from ammonium can occur. Clearly, atmospheric ammonium can contribute to soil acidification even at low soil pH.     

模拟氮沉降对温带典型森林土壤有效氮形态和含量的影响

通过室内模拟氮沉降试验,研究了氮沉降对温带典型森林土壤有效氮的影响.结果表明：试验期间,与对照相比,经过氮沉降处理的土壤铵态氮、硝态氮和有效氮均呈增长的趋势,增加的程度取决于森林类型、土层、氮处理类型和氮处理的持续时间.氮沉降对不同林型土壤有效氮形态和含量的影响不同,氮沉降对混交林的影响弱于阔叶林,强于针叶人工纯林;土壤A层对氮沉降的敏感程度大于土壤B层;铵态氮形态沉降对土壤铵态氮含量的影响比对土壤硝态氮含量的影响大,而硝态氮形态沉降对土壤硝态氮含量的影响比对土壤铵态氮含量的影响大,混合形态的氮沉降对二者均有促进作用,且增加幅度更高;氮沉降对土壤有效氮的影响存在累加效应.     

Nitrogen Oxide Fluxes and Nitrogen Cycling during Postagricultural Succession and Forest Fertilization in the Humid Tropics

The effects of changes in tropical land use on soil emissions of nitrous oxide (N₂O) and nitric oxide (NO) are not well understood. We examined emissions of N₂O and NO and their relationships to land use and forest composition, litterfall, soil nitrogen (N) pools and turnover, soil moisture, and patterns of carbon (C) cycling in a lower montane, subtropical wet region of Puerto Rico. Fluxes of N₂O and NO were measured monthly for over 1 year in old (more than 60 years old) pastures, early- and mid-successional forests previously in pasture, and late-successional forests not known to have been in pasture within the tabonuco (Dacryodes excelsa) forest zone. Additional, though less frequent, measures were also made in an experimentally fertilized tabonuco forest. N₂O fluxes exceeded NO fluxes at all sites, reflecting the consistently wet environment. The fertilized forest had the highest N oxide emissions (22.0 kg N · ha⁻¹· y⁻¹). Among the unfertilized sites, the expected pattern of increasing emissions with stand age did not occur in all cases. The mid-successional forest most dominated by leguminous trees had the highest emissions (9.0 kg N · ha⁻¹· y⁻¹), whereas the mid-successional forest lacking legumes had the lowest emissions (0.09 kg N · ha⁻¹· y⁻¹). N oxide fluxes from late-successional forests were higher than fluxes from pastures. Annual N oxide fluxes correlated positively to leaf litter N, net nitrification, potential nitrification, soil nitrate, and net N mineralization and negatively to leaf litter C:N ratio. Soil ammonium was not related to N oxide emissions. Forests with lower fluxes of N oxides had higher rates of C mineralization than sites with higher N oxide emissions. We conclude that (a) N oxide fluxes were substantial where the availability of inorganic N exceeded the requirements of competing biota; (b) species composition resulting from historical land use or varying successional dynamics played an important role in determining N availability; and (c) the established ecosystem models that predict N oxide loss from positive relationships with soil ammonium may need to be modified. Received 22 February 2000; accepted 6 September 2000.     

Watershed liming effects on the forest floor N cycle

The forest floor was expected to play a major role in determining the total ecosystem response to watershed liming because of its high concentration of nutrients and its high level of activity. Net N mineralization and net nitrification were estimated in a field survey using the buried-bag approach. In a laboratory incubation experiment, forest floor humus was mixed with 6 doses of lime to determine the sensitivity of N mineralization and nitrification to lime dose. Forest floor microcosms with and without live tree roots were used to calculate a N budget for the system.The pH of the forest floor increased from 3.6 to 4.9 in the Oe and to 4.0 in the Oa two years after liming. The extractable ammonium pool in both the field survey and microcosm study was substantially smaller after liming and was probably a result of the 36% to 55% lower net N mineralization rate in limed plots than in reference plots. The laboratory incubation results agreed with the field survey results and further demonstrated that at higher lime doses (pH 5 to 6), N mineralization increased above controls. Net nitrification in limed humus in both the buried bags and laboratory incubation was as much as three times higher than controls, which could explain why nitrate leaching in limed microcosms was greater than in control microcosms. However, nitrate leaching from microcosms with live. roots was not affected by liming, suggesting that roots in the forest floor may prevent excess nitrate leaching. Reductions in N mineralization had no effect on N leaching or N uptake, but reduced the extractable ammonium pool.     

Nitrogen cycling in an acid forest ecosystem in the Netherlands under increased atmospheric nitrogen input

Within a long-term research project studying the biogeochemical budget of an oak-beech forest ecosystem in the eastern part of the Netherlands, the nitrogen transformations and solute fluxes were determined in order to trace the fate of atmospherically deposited NH₄ ⁺ and to determine the contribution of nitrogen transformations to soil acidification.The oak-beech forest studied received an annual input of nitrogen via throughfall and stemflow of 45 kg N ha^–1 yr^–1, mainly as NH₄ ⁺, whereas 8 kg N ha^–1 yr^–1 was taken up by the canopy. Due to the specific hydrological regime resulting in periodically occurring high groundwater levels, denitrification was found to be the dominant output flux (35 kg N ha^–1 yr^–1). N₂0 emmission rate measurements indicated that 57% of this gaseous nitrogen loss (20 kg N ha^–1 yr^–1) was as N₂O. The forest lost an annual amount of 11 kg N ha^–1 yr^–1 via streamwater output, mainly as N0₃ ^–.Despite the acid conditions, high nitrification rates were measured. Nitrification occurred mainly in the litter layer and in the organic rich part of the mineral soil and was found to be closely correlated with soil temperature. The large amount of NH₄ ⁺ deposited on the forest floor via atmospheric deposition and produced by mineralization was to a large extent nitrified in the litter layer. Almost no NH₄ ⁺ reached the subsurface soil horizons. The N0₃ ^– was retained, taken up or transformed mainly in the mineral soil. A small amount of N0₃ ^– (9 kg N ha^–1 yr^–1) was removed from the system in streamwater output. A relatively small amount of nitrogen was measured in the soil water as Dissolved Organic Nitrogen.On the basis of these data the proton budget of the system was calculated using two different approaches. In both cases net proton production rates were high in the vegetation and in the litter layer of the forest ecosystem. Nitrogen transformations induced a net proton production rate of 2.4 kmol ha^–1 yr^–1 in the soil compartment.     

High precipitation causes large fluxes of dissolved organic carbon and nitrogen in a subtropical montane <Emphasis Type="Italic">Chamaecyparis</Emphasis> forest in Taiwan

Fluxes of dissolved organic carbon (DOC) and nitrogen (DON) may play an important role for losses of C and N from the soils of forest ecosystems, especially under conditions of high precipitation. We studied DOC and DON fluxes and concentrations in relation to precipitation intensity in a subtropical montane Chamaecyparis obtusa var. formosana forest in Taiwan. Our objective was, to quantify DOC and DON fluxes and to understand the role of high precipitation for DOC and DON export in this ecosystem. From 2005 to 2008 we sampled bulk precipitation, throughfall, forest floor percolates and seepage (60 cm) and analyzed DOC, DON and mineral N concentrations. Average DOC fluxes in the soil were extremely high (962 and 478 kg C ha^?1 year^?1 in forest floor percolates and seepage, respectively) while DON fluxes were similar to other (sub)tropical ecosystems (16 and 8 kg N ha^?1 year^?1, respectively). Total N fluxes in the soil were dominated by DON. Dissolved organic C and N concentrations in forest floor percolates were independent of the water flux. No dilution effect was visible. Instead, the pool size of potentially soluble DOC and DON was variable as indicated by different DOC and DON concentrations in forest floor percolates at similar precipitation amounts. Therefore, we hypothesized, that these pools are not likely to be depleted in the long term. The relationship between water fluxes in bulk precipitation and DOC and DON fluxes in forest floor percolates was positive (DOC r = 0.908, DON r = 0.842, respectively, Spearman rank correlation). We concluded, that precipitation is an important driver for DOC and DON losses from this subtropical montane forest and that these DOC losses play an important role in the soil C cycle of this ecosystem. Moreover, we found that the linear relationship between bulk precipitation and DOC and DON fluxes in forest floor percolates of temperate ecosystems does not hold when incorporating additional data on these fluxes from (subtropical) ecosystems.     

N uptake and N status in ponderosa pine as affected by soil compaction and forest floor removal

Soil compaction and forest floor removal influence fundamental soil processes that control forest productivity and sustainability. We investigated effects of soil compaction and forest floor removal on tree growth, N uptake and N status in ponderosa pine. Factorial combinations of soil compaction (non-compacted and compacted) and forest floor removal (forest floor present and no forest floor) were applied to three different surface soil textures. For studying N uptake, four trees from every treatment were ¹⁵N labeled with 130.6 mg m^–2 of ¹⁵N. Tree responses to compaction were dependent on the forest floor removal level. In loam and clay soils, non-compacted+no forest floor was beneficial to tree growth. Tree growth was depressed with compaction+no forest floor in clay soil. In sandy loam soil, compaction+no forest floor showed the best tree growth. No N deficiency was found in any soil type but a graphical method suggested correlation between N status and tree growth. In loam and clay soils, compaction+forest floor present increased N uptake. Nitrogen uptake was explained significantly by potential N mineralization in loam and clay soils. In sandy loam soil, the effects of compaction and forest floor removal were more complex, with the N uptake improved in the compaction+no forest floor treatment and reduced under compaction+forest floor present. Soil compaction may have influenced N tracer uptake because of improved unsaturated flow and root-soil contact. However, N immobilization may have restricted N uptake in compaction+forest floor present in the sandy loam soil. The study illustrates how soil properties and site preparation can potentially interact to affect N dynamics and forest productivity.     

A meta‐analysis of soil salinization effects on nitrogen pools,cycles and fluxes in coastal ecosystems

Salinity intrusion caused by land subsidence resulting from increasing groundwater abstraction, decreasing river sediment loads and increasing sea level because of climate change has caused widespread soil salinization in coastal ecosystems. Soil salinization may greatly alter nitrogen (N) cycling in coastal ecosystems. However, a comprehensive understanding of the effects of soil salinization on ecosystem N pools, cycling processes and fluxes is not available for coastal ecosystems. Therefore, we compiled data from 551 observations from 21 peer‐reviewed papers and conducted a meta‐analysis of experimental soil salinization effects on 19 variables related to N pools, cycling processes and fluxes in coastal ecosystems. Our results showed that the effects of soil salinization varied across different ecosystem types and salinity levels. Soil salinization increased plant N content (18%), soil NH₄⁺ (12%) and soil total N (210%), although it decreased soil NO₃^? (2%) and soil microbial biomass N (74%). Increasing soil salinity stimulated soil N₂O fluxes as well as hydrological NH₄⁺ and NO₂^? fluxes more than threefold, although it decreased the hydrological dissolved organic nitrogen (DON) flux (59%). Soil salinization also increased the net N mineralization by 70%, although salinization effects were not observed on the net nitrification, denitrification and dissimilatory nitrate reduction to ammonium in this meta‐analysis. Overall, this meta‐analysis improves our understanding of the responses of ecosystem N cycling to soil salinization, identifies knowledge gaps and highlights the urgent need for studies on the effects of soil salinization on coastal agro‐ecosystem and microbial N immobilization. Additional increases in knowledge are critical for designing sustainable adaptation measures to the predicted intrusion of salinity intrusion so that the productivity of coastal agro‐ecosystems can be maintained or improved and the N losses and pollution of the natural environment can be minimized.