Fluxes and concentrations of dissolved organic carbon and nitrogen – a synthesis for temperate forests

Dissolved organic carbon (DOC) and nitrogen (DON)represent an important part of the C and N cycles inforest ecosystems. Little is known about the controlson fluxes and concentrations of these compounds insoils under field conditions. Here we compiledpublished data on concentrations and fluxes of DOC andDON from 42 case studies in forest ecosystems of thetemperate zone in order to evaluate controls on alarger temporal and spatial scale. The focus was onannual fluxes and concentrations in throughfall,forest floor leachates and soil solutions. In allcompartments considered, concentrations and fluxesdiffered widely between the sites. Highestconcentrations of DOC and DON were generally observedin forest floor leachates and in A horizons. Highestfluxes occurred in forest floor leachates. The fluxesof DOC and DON in forest floor leachates increasedwith increasing annual precipitation and were alsopositively related to DOC and DON fluxes withthroughfall. Variation in throughfall fluxes couldexplain 46% and 65% of the variation in DOC and DONfluxes from the forest floor, respectively. No generaldifference in DOC and DON concentrations and fluxes inforest floor leachates was found when comparingconiferous and hardwood sites. Concentrations of DOCin forest floor leachates were positively correlatedto the pH of the forest floor. Furthermore, there wasno relationship between organic C and N stocks, soilC/N, litterfall or mineral N inputs and concentrationsand fluxes of DOC and DON in forest floor leachates.Including all compartments, fluxes of DOC and DON werehighly correlated. Ratios of DOC to DON calculatedfrom fluxes from the forest floor were independent ofthe amount of annual precipitation, pointing to asimilar response of DOC and DON to precipitationconditions. A decrease in the ratio of DOC to DON withsoil depth as observed on a plot-scale, was notconfirmed by data analysis on a large scale. Thecontrols observed on annual fluxes and concentrationsof DON and DOC at regional scale differed from thosereported for smaller time and space scales.     

Controls on the dynamics of dissolved organic carbon and nitrogen in a Central European deciduous forest

Despite growing attention concerning therole of dissolved organic matter (DOM) inelement cycling of forest ecosystems, thecontrols of concentrations and fluxes of bothdissolved organic carbon (DOC) and nitrogen(DON) under field conditions in forest soilsremain only poorly understood. The goal ofthis project is to measure the concentrations and fluxes of DON, NH₄ ⁺, NO₃ ^–and DOC in bulkprecipitation, throughfall, forest floorleachates and soil solutions of a deciduousstand in the Steigerwald region (northernBavaria, Germany). The DOC and DONconcentrations and fluxes were highest inleachates originating from the Oa layer of theforest floor (73 mg C L^–1, 2.3 mg NL^–1 and about 200–350 kg C, 8–10 kg Nha^–1 yr^–1). They were observed to behighly variable over time and decreased in themineral topsoil (17 mg C L^–1, 0.6 mg NL^–1 and about 50–90 kg C, 2.0 to 2.4 kg Nha^–1 yr^–1). The annual variability ofDOC and DON concentrations and subsequentialDOC/DON ratios was substantial in allsolutions. The DOC and DON concentrations inthroughfall were positively correlated withtemperature. The DOC and DON concentrationsdid not show seasonality in the forest floorand mineral soil. Concentrations were notrelated to litterfall dynamics but didcorrespond in part to the input of DOC and DONfrom throughfall. The throughfall contributionto the overall element fluxes was higher forDON than for DOC. Concentrations and fluxes ofDON were significantly correlated to DOC inthroughfall and the Oi layer. However, thecorrelation was weak in Oa leachates. Inaddition, seasonal and annual variation ofDOC/DON ratios indicated different mechanismsand release rates from the forest floor forboth components. The concentrations of DOC andDON in forest floor leachates were in mostcases dependent neither on the pH value orionic strength of the solution, nor on thewater flux or temperature changes. As aconsequence, the DOC and DON fluxes from theforest floor into the mineral soil werelargely dependent on the water flux if annualand biweekly time scales are considered.     

Concentration and Fluxes of Dissolved Organic Carbon (DOC) in Three Norway Spruce Stands along a Climatic Gradient in Sweden

Leaching of dissolved organic carbon (DOC) from the forest floor and transport in soil solution into the mineral soil are important for carbon cycling in boreal forest ecosystems. We examined DOC concentrations in bulk deposition, throughfall and in soil solutions collected under the O and B horizons in three Norway spruce stands along a climatic gradient in Sweden. Mean annual temperature for the three sites was 5.5, 3.4 and 1.2 °C. At each site we also examined the effect of soil moisture on DOC dynamics along a moisture gradient (dry, mesic and moist plots). To obtain information about the fate of DOC leached from the O horizon into the mineral soil, ¹⁴C measurements were made on bulk organic matter and DOC. The concentration and fluxes of DOC in O horizon leachates were highest at the southern site and lowest at the northern. Average DOC concentrations at the southern, central and northern sites were 49, 39 and 30 mg l⁻¹, respectively. We suggest that DOC leaching rates from O horizons were related to the net primary production of the ecosystem. Soil temperature probably governed the within-year variation in DOC concentration in O horizon leachates, but the peak in DOC was delayed relative to that of temperature, probably due to sorption processes. Neither soil moisture regime (dry, mesic or moist plots) nor seasonal variation in soil moisture seemed to be of any significance for the concentration of DOC leached from the O horizon. The ¹⁴C measurements showed that DOC in soil solution collected below the B horizon was derived mainly from the B horizon itself, rather than from the O horizon, indicating a substantial exchange (sorption–desorption reactions) between incoming DOC and soil organic carbon in the mineral soil.     

Leaching of dissolved organic carbon,dissolved organic nitrogen,and other solutes from coarse woody debris and litter in a mixed forest in New York State

Coarse woody debris (CWD) may play a role in nutrient cycling in temperate forests through the leaching of solutes, including dissolved organic carbon (DOC) and dissolved organic nitrogen (DON), to the underlying soil. These fluxes need to be considered in element budget calculations, and have the potential to influence microbial activity, soil development, and other processes in the underlying soil, but studies on leaching from CWD are rare. In this study, we collected throughfall, litter leachate, and CWD leachate in situ at a young mixed lowland forest in NY State, USA over one year. We measured the concentrations of DOC, DON, NH₄⁺, NO₃⁻, dissolved organic sulfur, SO₄²⁻, Cl⁻, Al, Ca, K, Mg, Na, and P, estimated the flux of these solutes in throughfall, and measured the cover of CWD to gain some insight into possible fluxes from CWD. Concentrations of DOC were much higher in CWD leachate than in throughfall or litter leachate (15 vs. 0.7 and 1.6 mM, respectively), and greater than reported values for other leachates from within forested ecosystems. Other solutes showed a similar pattern, with inorganic N being an exception. Our results suggest that microsite scale fluxes of DOC from CWD may be An high relative to throughfall and litter leaching fluxes, but since CWD covered a relatively small fraction (2%) of the forest floor in our study, ecosystem scale fluxes from CWD may be negligible for this site. Soil directly beneath CWD may be influenced by CWD leaching, in terms of soil organic matter, microbial activity, and N availability. Concentrations of some metals showed correlations to DOC concentrations, highlighting the possibility of complexation by DOM. Several solute concentrations in throughfall, including DOC, showed positive correlations to mean air temperature, and fewer showed positive correlations in litter leachate, while negative correlations were observed to precipitation, suggesting both biological and hydrologic control of solute concentrations.     

Dissolved organic carbon concentrations in four Norway spruce stands of different ages

Boreal forests are increasing in age partly due to reduced logging and efficient wildfire control. As a result, they also stock more carbon. Whether increased forest C stock causes greater production of dissolved organic carbon (DOC) is uncertain. DOC in bulk precipitation, throughfall and soil water was studied in 10-, 30-, 60- and 120-year-old stands of Norway spruce (Picea abies (L.) Karst.) DOC concentrations in throughfall and O horizon soil water followed the order 10 < 30 < 60 = 120 and 10 = 30 < 120 < 60, respectively. DOC fluxes followed the order 10 = 30 < 60 = 120 in throughfall, while no significant difference between stands was found for O horizon soil water. Above-ground tree litter varied according to 10 < 30 < 60 = 120, a pattern identical to that for DOC concentrations in throughfall and resembling but not identical to that for DOC concentrations in O horizon soil water. This indicates additional sources for DOC in soil water. Seasonality in DOC concentrations was observed at the base of the O horizon, and seasonality in DOC fluxes in both throughfall and O horizon soil water. Our results suggest differences in the polarity of DOC between the 10-year stand and the others, which we interpret as reflecting the lack of grown trees and possibly the different vegetation on the 10-year stand.     

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.     

The effects of gap disturbance on nitrogen cycling and retention in late-successional northern hardwood–hemlock forests

Late-successional forests in the upper Great Lakes region are susceptible to nitrogen (N) saturation and subsequent nitrate (NO₃⁻) leaching loss. Endemic wind disturbances (i.e., treefall gaps) alter tree uptake and soil N dynamics; and, gaps are particular susceptible to NO₃⁻ leaching loss. Inorganic N was measured throughout two snow-free periods in throughfall, forest floor leachates, and mineral soil leachates in gaps (300–2,000 m², 6–9 years old), gap-edges, and closed forest plots in late-successional northern hardwood, hemlock, and northern hardwood–hemlock stands. Differences in forest water inorganic N among gaps, edges, and closed forest plots were consistent across these cover types: NO₃⁻ inputs in throughfall were significantly greater in undisturbed forest plots compared with gaps and edges; forest floor leachate NO₃⁻ was significantly greater in gaps compared to edges and closed forest plots; and soil leachate NO₃⁻ was significantly greater in gaps compared to the closed forest. Significant differences in forest water ammonium and pH were not detected. Compared to suspected N-saturated forests with high soil NO₃⁻ leaching, undisturbed forest plots in these late-successional forests are not losing NO₃⁻ (net annual gain of 2.8 kg ha⁻¹) and are likely not N-saturated. Net annual NO₃⁻ losses were observed in gaps (1.3 kg ha⁻¹) and gap-edges (0.2 kg ha⁻¹), but we suspect these N leaching losses are a result of decreased plant uptake and increased soil N mineralization associated with disturbance, and not N-saturation.     

Vertical transport of dissolved organic C and N under long-term N amendments in pine and hardwood forests

At the Harvard Forest, Massachusetts, a long-term effort is under way to study responses in ecosystem biogeochemistry to chronic inputs of N in atmospheric deposition in the region. Since 1988, experimental additions of NH₄NO₃ (0, 5 and 15 g N m^–2 yr^–1) have been made in two forest stands:Pinus resinosa (red pine) and mixed hardwood. In the seventh year of the study, we measured solute concentrations and estimated solute fluxes in throughfall and at two soil depths, beneath the forest floors (Oa) and beneath the B horizons.Beneath the Oa, concentrations and fluxes of dissolved organic C and N (DOC and DON) were higher in the coniferous stand than in the hardwood stand. The mineral soil exerted a strong homogenizing effect on concentrations beneath the B horizons. In reference plots (no N additions), DON composed 56% (pine) and 67% (hardwood) of the total dissolved nitrogen (TDN) transported downward from the forest floor to the mineral soil, and 98% of the TDN exported from the solums. Under N amendments, fluxes of DON from the forest floor correlated positively with rates of N addition, but fluxes of inorganic N from the Oa exceeded those of DON. Export of DON from the solums appeared unaffected by 7 years of N amendments, but as in the Oa, DON composed smaller fractions of TDN exports under N amendments. DOC fluxes were not strongly related to N amendment rates, but ratios of DOC:DON often decreased.The hardwood forest floor exhibited a much stronger sink for inorganic N than did the pine forest floor, making the inputs of dissolved N to mineral soil much greater in the pine stand. Under the high-N treatment, exports of inorganic N from the solum of the pine stand were increased >500-fold over reference (5.2 vs. 0.01 g N m^–2 yr^–1), consistent with other manifestations of nitrogen saturation. Exports of N from the solum in the pine forest decreased in the order NO₃-N> NH₄-N> DON, with exports of inorganic N 14-fold higher than exports of DON. In the hardwood forest, in contrast, increased sinks for inorganic N under N amendments resulted in exports of inorganic N that remained lower than DON exports in N-amended plots as well as the reference plot.     

Fluorescence characteristics and biodegradability of dissolved organic matter in forest and wetland soils from coastal temperate watersheds in southeast Alaska

Understanding how the concentration and chemical quality of dissolved organic matter (DOM) varies in soils is critical because DOM influences an array of biological, chemical, and physical processes. We used PARAFAC modeling of excitation–emission fluorescence spectroscopy, specific UV absorbance (SUVA₂₅₄) and biodegradable dissolved organic carbon (BDOC) incubations to investigate the chemical quality of DOM in soil water collected from 25 cm piezometers in four different wetland and forest soils: bog, forested wetland, fen and upland forest. There were significant differences in soil solution concentrations of dissolved organic C, N, and P, DOC:DON ratios, SUVA₂₅₄ and BDOC among the four soil types. Throughout the sampling period, average DOC concentrations in the four soil types ranged from 9–32 mg C l⁻¹ and between 23–42% of the DOC was biodegradable. Seasonal patterns in dissolved nutrient concentrations and BDOC were observed in the three wetland types suggesting strong biotic controls over DOM concentrations in wetland soils. PARAFAC modeling of excitation–emission fluorescence spectroscopy showed that protein-like fluorescence was positively correlated (r ² = 0.82; P < 0.001) with BDOC for all soil types taken together. This finding indicates that PARAFAC modeling may substantially improve the ability to predict BDOC in natural environments. Coincident measurements of DOM concentrations, BDOC and PARAFAC modeling confirmed that the four soil types contain DOM with distinct chemical properties and have unique fluorescent fingerprints. DOM inputs to streams from the four soil types therefore have the potential to alter stream biogeochemical processes differently by influencing temporal patterns in stream heterotrophic productivity.     

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.     

Site‐to‐site variability and temporal trends of DOC concentrations and fluxes in temperate forest soils

Here, we report site‐to‐site variability and 12–14 year trends of dissolved organic carbon (DOC) from organic layers and mineral soils of 22 forests in Bavaria, Germany. DOC concentrations in the organic layer were negatively correlated with mean annual precipitation and elevation whereas air temperature had a positive effect on DOC concentrations. DOC fluxes in subsoils increased by 3 kg ha^?1 yr^?1 per 100 mm precipitation or per 100 m elevation. The highest DOC concentrations were found under pine stands with mor humus. Average DOC concentrations in organic layer leachates followed the order: pine>oak>spruce>beech. However, the order was different for mean DOC fluxes (spruce>pine>oak>beech) because of varying precipitation regimes among the forest types. In 12 of 22 sites, DOC concentrations of organic layer leachates significantly increased by 0.5 to 3.1 mg C L^?1 yr^?1 during the sampling period. The increase in DOC concentration coincided with decreasing sulfate concentration, indicating that sulfate concentration is an important driver of DOC solubility in the organic layer of these forest sites. In contrast to the organic layer, DOC concentrations below 60 cm mineral soil depth decreased by <0.1–0.4 mg C L^?1 yr^?1 at eight sites. The negative DOC trends were attributed to (i) increasing adsorption of DOC by mineral surfaces resulting from desorption of sulfate and (ii) increasing decay of DOC resulting from decreasing stabilization of DOC by organo‐Al complexes. Trends of DOC fluxes from organic layers were consistent with those of DOC concentrations although trends were only significant at seven sites. DOC fluxes in the subsoil were with few exceptions small and trends were generally not significant. Our results suggest that enhanced mobilization of DOC in forest floors contributed to the increase of DOC in surface waters while mineral horizons did not contribute to increasing DOC export of forest soils.     

Dynamics of dissolved organic <Superscript>14</Superscript>C in throughfall and soil solution of a Norway spruce forest

Dissolved organic carbon (DOC) is an important component of the C cycle in forest ecosystems, but dynamics and origin of DOC in throughfall and soil solution are yet poorly understood. In a 2-year study, we analyzed the radiocarbon signature of DOC in throughfall and soil solution beneath the Oa horizon and at 90 cm depth in a Norway spruce forest on a Podzol soil. A two-pool mixing model revealed that throughfall DOC comprised mainly biogenic C, i.e. recently fixed C, from canopy leaching and possibly other sources. The contribution of fossil DOC from atmospheric deposition to throughfall DOC was on average 6% with maxima of 8–11% during the dormant season. In soil solution from the Oa horizon, DO¹⁴C signature was highly dynamic (range from −8‰ to +103‰), but not correlated with DOC concentration. Radiocarbon signatures suggest that DOC beneath the Oa horizon originated mainly from occluded and mineral associated organic matter fractions of the Oa horizon rather than from the Oi or Oe horizon. Relatively old C was released in the rewetting phase following a drought period in the late summer of 2006. In contrast, the DO¹⁴C signature indicated the release of younger C throughout the humid year 2007. In soil solutions from 90 cm depth, DO¹⁴C signatures were also highly dynamic (−127‰ to +3‰) despite constantly low DOC concentrations. Similar to the Oa horizon, the lowest DO¹⁴C signature at 90 cm depth was found after the rewetting phase in the late summer of 2006. Because of the variation in the DO¹⁴C signatures at this depth, we conclude that DOC was not equilibrated with the surrounding soil, but also originated from overlaying soil horizons. The dynamics of DO¹⁴C in throughfall and soil solution suggest that the sources of DOC are highly variable in time. Extended drought periods likely have a strong influence on release and translocation of DOC from relatively old and possibly stabilized soil organic matter fractions. Temporal variations as well as the input of fossil DOC needs to be considered when calibrating DOC models based on DO¹⁴C signatures.     

Characterization of solid and dissolved carbon in a spruce-fir Spodosol

Organic substances are an integral part of the biogeochemistry of many elements in forest ecosystems. However, our understanding of the composition, chemistry, and reactions of these materials are incomplete and sometimes inconsistent. Therefore, we examined in detail dissolved organic carbon (DOC) in forest floor leachates over a two-year period (1992–1993), soil C, and DOC adsorption by a mineral soil to determine the relationship between soil solid and solution C characteristics in a spruce-fir ecosystem. The structural composition of DOC, DOC fractions (hydrophobic and hydrophilic acids, hydrophilic neutrals), and soil samples from the organic and mineral horizons were also analyzed using¹³C nuclear magnetic resonance (NMR) spectroscopy.Total DOC in forest floor leachates ranged from 7.8 to 13.8 mmol L^–1 with an average of 8.6 mmol L^–1. Concentrations were highest in September of both 1992 and 1993. Fractionation of the forest floor DOC indicated these solutions contained high organic acid contents that averaged 92% of the total DOC. Hydrophobic acids were also preferentially adsorbed by the B horizon. The¹³C NMR data suggested alkyl, carbohydrate, aromatic, and carboxylic C were the primary constituents for organic and mineral soils, DOC, and DOC fractions. Compositional changes of C were observed as aromatic and carbohydrate decreased, whereas alkyl, methoxy, and carbonyl moieties increased with depth. However, C composition changed little among the three organic layers based on the similarity of alkyl/carbohydrates ratios as determined from NMR area integration, suggesting that in this acid soil, decomposition proceeds rather slowly. Hydrophobic acids contained high contents of aromatic C, whereas hydrophilic acids were comprised primarily of carboxylic C. Hydrophilic neutrals were rich in carbohydrate C. Results indicated that these DOC fractions were unaltered during the isolation process. Carboxylic C groups appeared to dissolve easily and were probably the primary contributor to organic acidity in our organic dominate leachates. Results also suggested that DOC materials adsorbed on the B horizon underwent further biodegradation. Several seasonal patterns of C composition were observed in the forest floor leachates and DOC fractions collected between 1992 and 1993.Overall, the evidence from this study suggested that (i) DOC levels were mainly controlled by biological activity, (ii) forest floor DOC was comprised primarily of organic acids, (iii) contact of soil leachates with B horizon material affected DOC quantitatively and qualitatively, (iv) phenolic, carboxylic, and carbonyl C appeared to dissolve readily in the forest Oa horizon, (v) DOC materials adsorbed on the B horizon selectively underwent further decomposition, and (vi) C composition is a function of the extent of decomposition and DOC fractions.     

Controls on the release of dissolved organic carbon and nitrogen from a deciduous forest floor investigated by manipulations of aboveground litter inputs and water flux

Although dissolved organic matter (DOM) released from the forest floor plays a crucial role in transporting carbon and major nutrients through the soil profile, its formation and responses to changing litter inputs are only partially understood. To gain insights into the controlling mechanisms of DOM release from the forest floor, we investigated responses of the concentrations and fluxes of dissolved organic carbon (DOC) and nitrogen (DON) in forest floor leachates to manipulations of throughfall (TF) flow and aboveground litter inputs (litter removal, litter addition, and glucose addition) at a hardwood stand in Bavaria, Germany. Over the two-year study period, litter manipulations resulted in significant changes in C and N stocks of the uppermost organic horizon (Oi). DOC and DON losses via forest floor leaching represented 8 and 11% of annual litterfall C and N inputs at the control, respectively. The exclusion of aboveground litter inputs caused a slight decrease in DOC release from the Oi horizon but no change in the overall leaching losses of DOC and DON in forest floor leachates. In contrast, the addition of litter or glucose increased the release of DOC and DON either from the Oi or from the lower horizons (Oe + Oa). Net releases of DOC from the Oe + Oa horizons over the entire manipulation period were not related to changes in microbial activity (measured as rates of basal and substrate-induced respiration) but to the original forest floor depths prior to manipulation, pointing to the flux control by the size of source pools rather than a straightforward relationship between microbial activity and DOM production. In response to doubled TF fluxes, net increases in DOM fluxes occurred in the lower forest floor, indicating the presence of substantial pools of potentially soluble organic matter in the Oe + Oa horizons. In contrast to the general assumption of DOM as a leaching product from recent litter, our results suggest that DOM in forest floor leachates is derived from both newly added litter and older organic horizons through complex interactions between microbial production and consumption and hydrologic transport.     

Rainfall partitioning and related hydrochemical fluxes in a diverse and in a mono specific (Phenakospermum guyannense) secondary vegetation stand in eastern Amazonia

Rainfall partitioning into throughfall and stemflow was studied in a diverse and in a mono specific stand of secondary vegetation in Eastern Amazonia. The nutrient concentrations in the water were analysed in order to quantify the related hydrochemical fluxes. Secondary vegetation forms the fallow in the local shifting cultivation system and is usually dominated by shrubs and trees. Phenakospermum guyannense (Strelitziaceae), a banana-like herb, is one of the predominant non-woody species. The study was conducted during an 18-month period in a 2.5-year-old relatively species-rich stand and a 10-year-old stand dominated by P. guyannense. In a year with 1956 mm of rainfall 65% (1281 mm) of this quantity reached the soil as throughfall in the diverse stand and 38% (743 mm) in the mono specific stand. Stemflow was estimated to be 23% and 41% respectively. P. guyannense and Banara guianensis (Flacourtiaceae), a tree species, were causing these high funnelling effects. In the young diverse stand B. guianensis had a stemflow of more than 200 l year⁻¹ and P. guyannense had a median flux of 77 l year⁻¹ per pseudostem. In the older stand the taller plants of P.␣guyannense collected 644 l year⁻¹ per pseudostem on the median. The reason for these high values could be the banana-like growth form of P. guyannense and the crown morphology of B. guianensis, which has inclined branches. The low proportion of throughfall and the high stemflow values differ from all previous studies in Amazonian primary forests. The proximity to the Atlantic Ocean strongly influenced the nutrient fluxes via rainfall at our study site. This becomes obvious from the high Na and Cl fluxes with rainfall (19.7 kg Na ha⁻¹ year⁻¹, 37.2 kg Cl ha⁻¹ year⁻¹) which were approximately equal to the Na and Cl fluxes with the sum of throughfall and stemflow for both stands. K fluxes in throughfall and stemflow in both stands were higher than in rainfall by a factor of 8. The high K enrichment during the crown passage is assumed to be caused by a␣high K concentration in the leaf tissue resulting in enhanced leaching from the leaves. In months with low␣rainfall the concentrations of Ca, Mg, S and Cl in throughfall of the diverse stand were significantly higher than in months with high rainfall. This was mainly due to vegetation burns in the dry period, which resulted in ash deposition on the canopy and subsequent wash-off and solution of ash particles. Received: 11 May 1997 / Accepted: 2 November 1997     

Carbon and nitrogen storage in an age-sequence of <Emphasis Type="Italic">Pinus densiflora</Emphasis> stands in Korea

The carbon (C) and nitrogen (N) storage capabilities of Pinus densiflora in six different stand ages (10, 27, 30, 32, 44, and 71 years old) were investigated in Korea. Thirty sample trees were destructively harvested and 12 were excavated. Samples from the above and belowground tree components, coarse woody debris (CWD), forest floor, and mineral soil (0–30 cm) were collected. Tree biomass was highest in the 71-year-old stand (202.8 t ha⁻¹) and lowest in the 10-year-old stand (18.4 t ha⁻¹). C and N storage in the mineral soil was higher in the 71-year-old stand than in the other stands, mainly due to higher soil C and N concentrations. Consequently, the total ecosystem C and N storage (tree+forest floor+CWD+soil) was positively correlated with stand age: increasing from a minimum in the 10 year old stand (18.8 t C ha⁻¹ and 1.3 t N ha⁻¹) to a maximum in the 71-year-old stand (201.4 t C ha⁻¹ and 8.5 t N ha⁻¹). The total ecosystem C storage showed a similar sigmoidal pattern to that of tree C storage as a function of the age-sequence, while N storage in the CWD, forest floor and mineral soil showed no significant temporal trends. Our results provide important insights that will increase our understanding of C and N storage in P. densiflora stands and our ability to predict changes according to stand age in the region.     

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.     

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.     

Nitrogen Cycling and Mass Balance for a Forested Catchment in the Italian Alps. Assessment of Nitrogen Status

During 1999–2001 the chemical composition and fluxes were measured in rainfall, throughfall, soil solution and stream water in a remote forested site in the Italian Alps. The analysis of temporal patterns revealed the differential behaviour of nitrogen and sulphur and suggested that different mechanisms controlled their flux. No important changes in sulphate concentration and fluxes emerged as the solution passed through the various components of the forest ecosystem, and temporal variations of SO₄ in the soil solution and stream were likely driven by the physical process of dilution. The availability of nitrate and ammonia, by contrast, was drastically reduced as throughfall water entered the soil and passed through the mineral layers, irrespective of season. The calculated hydrochemical budget based on throughfall and soil solution N fluxes revealed that ~80% N retention in the forest soil, corresponding to 12 kg ha⁻¹ yr⁻¹, despite a relatively high N deposition loading (15 kg ha⁻¹ yr⁻¹). Most of the leached nitrogen (90%) was in the organic form. Indicators of the N status of this ecosystem, such as C/N ratio in solid and solution phase of the soil and N foliage content as well as land use history were examined. Despite the strong N retention in the forested part of the catchment, the stream water N–NO₃ levels were consistently above 10 μg l⁻¹ suggesting that the Val Masino catchment as a whole was less efficient in processing atmospheric N inputs. This contrasting N behaviour illustrates the role of landscape features, such as the soil cover and vegetation type, that is characteristic of an alpine catchment.     

Litterfall and fine root biomass contribution to nutrient dynamics in second- and old-growth Douglas-fir ecosystems

Litterfall and fine root production were measured for three years as part of a carbon balance study of three forest stands in the Pacific Northwest of the United States. A young second-growth Douglas-fir [Pseudotsuga menziesii (Mirb.) Franco] stand, a second-growth Douglas-fir with red alder (Alnus rubra Bong.) stand, and an old-growth (∼550 years) Douglas-fir stand were monitored for inputs of carbon and nitrogen into the soil from litterfall and fine root production, as well as changes in soil C and N. Fine root production and soil nutrient changes were measured through the use of soil ingrowth bags containing homogenized soil from the respective stands. Litterfall biomass was greatest in the Douglas-fir-alder stand (527 g m⁻² yr⁻¹) that annually returned nearly three times the amount of N as the other stands. Mean residence time for forest floor material was also shortest at this site averaging 4.6 years and 5.5 years for C an N, respectively. Fine root production in the upper 20 cm ranged from 584 g m⁻² in the N rich Douglas-fir-alder stand to 836 g m⁻² in the old-growth stand. Fine root production (down to one meter) was always greater than litterfall with a below:above ratio ranging from 3.73 for the young Douglas-fir stand to 1.62 for the Douglas-fir-alder stand. The below:above N ratios for all three stands closely approximate those for biomass. Soil changes in both C and N differed by site, but the soil C changes in the old-growth stand mirrored those obtained in an ongoing CO₂ flux study. Results from the soil ingrowth bags strongly suggest that this method provides a simple, but sufficient device for measuring potential fine root biomass production as well as soil chemical changes.