Decomposition pathways of 13C-depleted leaf litter in forest soils of the Swiss Jura

Decomposition of leaf litter and its incorporation into the mineral soil are key components of the C cycle in forest soils. In a ¹³C tracer experiment, we quantified the pathways of C from decomposing leaf litter in calcareous soils of a mixed beech forest in the Swiss Jura. Moreover, we assessed how important the cold season is for the decomposition of freshly fallen leaves. The annual C loss from the litter layer of 69–77% resulted mainly from the C mineralization (29–34% of the initial litter C) and from the transfer of litter material to the deeper mineral soil (>4 cm) by soil fauna (30%). Although only 4–5% of the initial litter C was leached as dissolved organic carbon (DOC), this pathway could be important for the C sequestration in soils in the long term: The DOC leached from the litter layer was mostly retained (95%) in the first 5 cm of the mineral soil by both physico-chemical sorption and biodegradation and, thus, it might have contributed significantly to the litter-derived C recovered in the heavy fraction (>1.6 g cm^?3) at 0–4 cm depth (4% of the initial litter C). About 80% of the annual DOC leaching from the litter layer occurred during the cold season (Nov–April) due to an initial DOC flush of water-soluble substances. In contrast, the litter mineralization in winter accounted for only 25% of the annual C losses through CO₂ release from the labelled litter. Nevertheless, the highest contributions (45–60%) of litter decay to the heterotrophic soil respiration were observed on warm winter days when the mineral soil was still cold and the labile litter pool only partly mineralized. Our ¹³C tracing also revealed that: (1) the fresh litter C only marginally primed the mineralization of older SOM (>1 year); and (2) non-litter C, such as throughfall DOC, contributed significantly to the C fluxes from the litter layer since the microbial biomass and the DOC leached from the litter layer contained 20–30% and up to 60% of unlabelled C, respectively. In summary, our study shows that significant amounts of recent leaf litter C (<1 year) are incorporated into mineral soils and that the cold season is clearly less important for the litter turnover than the warm season in this beech forest ecosystem.     

Nitrogen addition alters mineralization dynamics of 13C‐depleted leaf and twig litter and reduces leaching of older DOC from mineral soil

Recent reviews indicate that N deposition increases soil organic matter (SOM) storage in forests but the undelying processes are poorly understood. Our aim was to quantify the impacts of increased N inputs on soil C fluxes such as C mineralization and leaching of dissolved organic carbon (DOC) from different litter materials and native SOM. We added 5.5 g N m^?2 yr^?1 as NH₄NO₃ over 1 year to two beech forest stands on calcareous soils in the Swiss Jura. We replaced the native litter layer with ¹³C‐depleted twigs and leaves (δ¹³C: ?38.4 and ?40.8‰) in late fall and measured N effects on litter‐ and SOM‐derived C fluxes. Nitrogen addition did not significantly affect annual C losses through mineralization, but altered the temporal dynamics in litter mineralization: increased N inputs stimulated initial mineralization during winter (leaves: +25%; twigs: +22%), but suppressed rates in the subsequent summer. The switch from a positive to a negative response occurred earlier and more strongly for leaves than for twigs (?21% vs. 0%). Nitrogen addition did not influence microbial respiration from the nonlabeled calcareous mineral soil below the litter which contrasts with recent meta‐analysis primarily based on acidic soils. Leaching of DOC from the litter layer was not affected by NH₄NO₃ additions, but DOC fluxes from the mineral soils at 5 and 10 cm depth were significantly reduced by 17%. The ¹³C tracking indicated that litter‐derived C contributed less than 15% of the DOC flux from the mineral soil, with N additions not affecting this fraction. Hence, the suppressed DOC fluxes from the mineral soil at higher N inputs can be attributed to reduced mobilization of nonlitter derived ‘older’ DOC. We relate this decline to an altered solute chemistry by NH₄NO₃ additions, an increased ionic strength and acidification resulting from nitrification, rather than to a change in microbial decomposition.     

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.     

Dissolved organic matter under native Cerrado and Pinus caribaea plantations in the Brazilian savanna

The transformation of native Cerrado into Pinus caribaea Morelet plantations changes the DOM dynamics including changed rates of mineralisation, denitrification, and C export to the groundwater. To examine the differences in quantity, temporal dynamics, and quality of DOM between Cerrado and Pine plantations we collected rainfall, throughfall, stemflow, litter leachate (under pine only) and soil solution at 15, 80, and 200 cm depth in weekly intervals during the rainy seasons 1997/98 and 1998/99. We determined total dissolved organic carbon (DOC) concentrations and assessed DOM quality by separating hydrophilic and hydrophobic fractions and by NMR analysis of organic layer extracts. The rainfall had a mean DOC concentration of 2.6 mg L^–1. The mean concentrations of DOC in the throughfall of the pine plantations (5.0–10.5 mg L^–1) were significantly above those of Cerrado (2.6–4.9 mg L^–1). During the first part of the rainy seasons (October–December), the concentrations of DOC in the soil solution (15–200 cm depth) under Cerrado and pine did not differ significantly. During the second part of both rainy seasons (January–April), the concentrations of DOC in the soil solution under Cerrado (4.4–5.1 mg L^–1) exceeded those under PI (1.4–2.7 mg L^–1). Possible explanations of the latter include higher DOM input into the Cerrado soil and a stronger retention and/or faster mineralisation of the pine DOM than of the Cerrado DOM in the mineral soil. As the structural composition of DOM extracted from the organic layer under Cerrado and pine did not differ significantly, faster mineralisation was the most likely explanation for partly lower DOC concentrations in the soil solution under pine than under Cerrado. This assumption was supported by increasing contributions of hydrophobic DOM to total DOM with increasing depth under pine while, under Cerrado, the DOM composition did not change with depth. The reason for DOM mineralisation under pine was probably the higher N availability because total N concentrations were 11–23 times higher under pine than under Cerrado.     

Composition,Dynamics, and Fate of Leached Dissolved Organic Matter in Terrestrial Ecosystems: Results from a Decomposition Experiment

Fluxes of dissolved organic matter (DOM) are an important vector for the movement of carbon (C) and nutrients both within and between ecosystems. However, although DOM fluxes from throughfall and through litterfall can be large, little is known about the fate of DOM leached from plant canopies, or from the litter layer into the soil horizon. In this study, our objectives were to determine the importance of plant-litter leachate as a vehicle for DOM movement, and to track DOM decomposition [including dissolve organic carbon (DOC) and dissolved organic nitrogen (DON) fractions], as well as DOM chemical and isotopic dynamics, during a long-term laboratory incubation experiment using fresh leaves and litter from several ecosystem types. The water-extractable fraction of organic C was high for all five plant species, as was the biodegradable fraction; in most cases, more than 70% of the initial DOM was decomposed in the first 10 days of the experiment. The chemical composition of the DOM changed as decomposition proceeded, with humic (hydrophobic) fractions becoming relatively more abundant than nonhumic (hydrophilic) fractions over time. However, in spite of proportional changes in humic and nonhumic fractions over time, our data suggest that both fractions are readily decomposed in the absence of physicochemical reactions with soil surfaces. Our data also showed no changes in the ¹³C signature of DOM during decomposition, suggesting that isotopic fractionation during DOM uptake is not a significant process. These results suggest that soil microorganisms preferentially decompose more labile organic molecules in the DOM pool, which also tend to be isotopically heavier than more recalcitrant DOM fractions. We believe that the interaction between DOM decomposition dynamics and soil sorption processes contribute to the ¹³C enrichment of soil organic matter commonly observed with depth in soil profiles. published online 2004     

Detrital Controls on Soil Solution N and Dissolved Organic Matter in Soils: A Field Experiment

We established a long-term field study in an old growth coniferous forest at the H.J. Andrews Experimental Forest, OR, USA, to address how detrital quality and quantity control soil organic matter accumulation and stabilization. The Detritus Input and Removal Treatments (DIRT) plots consist of treatments that double leaf litter, double woody debris inputs, exclude litter inputs, or remove root inputs via trenching. We measured changes in soil solution chemistry with depth, and conducted long-term incubations of bulk soils from different treatments in order to elucidate effects of detrital inputs on the relative amounts and lability of different soil C pools. In the field, the addition of woody debris increased dissolved organic carbon (DOC) concentrations in O-horizon leachate and at 30 cm, but not at 100 cm, compared to control plots, suggesting increased rates of DOC retention with added woody debris. DOC concentrations decreased through the soil profile in all plots to a greater degree than did dissolved organic nitrogen (DON), most likely due to preferential sorption of high C:N hydrophobic dissolved organic matter (DOM) in upper horizons; percent hydrophobic DOM decreased significantly with depth, and hydrophilic DOM had a much lower and less variable C:N ratio. Although laboratory extracts of different litter types showed differences in DOM chemistry, percent hydrophobic DOM did not differ among soil solutions from different detrital treatments in the field, suggesting that microbial processing of DOM leachate in the field consumed easily degradable components, thus equalizing leachate chemistry among treatments. Total dissolved N leaching from plots with intact roots was very low (0.17 g m⁻² year⁻¹), slightly less than measured deposition to this very unpolluted forest (~s 0.2 g m⁻² year⁻¹). Total dissolved N losses showed significant increases in the two treatments without roots whereas concentrations of DOC decreased. In these plots, N losses were less than half of estimated plant uptake, suggesting that other mechanisms, such as increased microbial immobilization of N, accounted for retention of N in deep soils. In long-term laboratory incubations, soils from plots that had both above- and below-ground litter inputs excluded for 5 years showed a trend towards lower DOC loss rates, but not lower respiration rates. Soils from plots with added wood had similar respiration and DOC loss rates as control soils, suggesting that the additional DOC sorption observed in the field in these soils was stabilized in the soil and not readily lost upon incubation.     

Initial differentiation of vertical soil organic matter distribution and composition under juvenile beech (Fagus sylvatica L.) trees

In a lysimeter experiment with juvenile beech trees (Fagus sylvatica L.) we studied the development of depth gradients of soil organic matter (SOM) composition and distribution after soil disturbance. The sampling scheme applied to the given soil layers (0–2 cm, 2–5 cm, 5–10 cm and 10–20 cm) was crucial to study the subtle reformation of SOM properties with depth in the artificially filled lysimeters. Due to the combination of physical SOM fractionation with the application of ¹⁵N-labelled beech litter and ¹³C-CPMAS NMR spectroscopy we were able to obtain a detailed view on vertical differentiation of SOM properties. Four years after soil disturbance a significant decrease of the mass of particulate OM (POM) with depth could be found. A clear depth distribution was also shown for carbon (C) and nitrogen (N) within the SOM fractions related to bulk soil. The mineral fractions <63 µm clearly dominated C storage (between 47 to 60% of bulk soil C) and N storage (between 68 to 86% of bulk soil N). A drastic increase in aliphatic C structures concomitant to decreasing O/N-alkyl C was detected with depth, increasing from free POM to occluded POM. Only a slight depth gradient was observed for ¹³C but a clear vertical incorporation of ¹⁵N from the applied labelled beech litter was demonstrated probably resulting from faunal and fungal incorporation. We clearly demonstrated a significant reformation of a SOM depth profile within a very short time of soil evolution. One important finding of this study is that especially in soils with reforming SOM depth gradients after land-use changes selective sampling of whole soil horizons can bias predictions of C and N dynamics as it overlooks a potential development of gradients of SOM properties on smaller scales.     

氮沉降对杉木人工林土壤可溶性有机质数量和结构的影响

氮沉降作为现在乃至未来气候变化的趋势之一,其可能深刻影响土壤可溶性有机质的数量和结构。选取我国中亚热带杉木人工林不同深度土壤（0-10 cm和10-20 cm）进行氮沉降试验,利用光谱技术研究氮沉降对土壤可溶性有机质数量和结构的影响。试验设对照（CT,0 kg hm^-2 a^-1）、高氮（HN：80 kg hm^-2 a^-1）、低氮（LN：40 kg hm^-2 a^-1）3种处理。结果表明：（1）在0-10 cm和10-20 cm土层,HN、LN处理的土壤可溶性有机碳和可溶性有机氮含量显著高于CT。（2）在0-10 cm和10-20 cm土层,1月时HN、LN处理的芳香性指数和腐殖化程度都显著高于CT,而4月时HN、LN处理的芳香性指数和腐殖化程度都显著低于CT。除了氮含量的直接影响外,RDA （冗余分析）表明,两土层中土壤含水量、pH和土壤有机碳是驱动氮沉降对土壤DOM数量和结构的重要环境因子。因此,氮沉降对土壤DOM的影响是复杂的,未来尤其应该注重氮沉降对生态系统影响的季节模式。     

Decomposition of roots and twigs: Effects of wood type (beech and ash), diameter,site of exposure and macrofauna exclusion

Carbon loss and nitrogen dynamics in beech roots (Fagus sylvatica L.), beech twigs and ash roots (Fraxinus excelsior L.) of 0–3, 3–10 and 10–40 mm diameter were investigated during 36 months of exposure in litter bags of 1 and 4 mm mesh. Four experiments were set up: (1) Beech and ash roots (three size classes) were placed in a soil depth of ca 5 cm in a beechwood on limestone; (2) beech twigs (three size classes) were placed on the soil surface of the beechwood; (3) beech roots (3–10 mm) were placed on the soil surface of the beechwood: (4) beech twigs (3–10 mm) were placed on the soil surface of four sites representing different stages of secondary succession (wheat field, 13 year old fallow, ca 50 year old fallow, beechwood). Ash roots generally lost more C than beech roots. Loss in C of ash roots was similar for each of the size classes, whereas in beech roots and beech twigs C loss was in the order large roots > medium roots > small roots. Beech roots (3–10 mm) placed on the soil surface lost considerably less C than beech twigs (3–10 mm). Decomposition of beech twigs varied among ecosystems but generally did not follow clear patterns with successional stages. The fit of linear vs exponential models of decay is compared and in most materials exponential models fitted the data better. In each of the wood materials an accumulation of N occurred. Irrespective of wood type, root and twig diameter, mineralization of N of wood materials placed in the beechwood started uniformly after 12 months. Multiple regression analysis indicated a negative relationship between initial N content and C loss in beech roots and twigs but not in ash roots. The analysis also indicated a significant influence of the degree of white rot and of the amount of mineral soil deposited in the litter bags on C loss of certain wood materials. Generally, mesh size affected C loss and N dynamics only slightly, which is attributed to the comparatively short exposure time.     

An assessment,using stable isotopes,of the importance of allochthonous organic carbon sources to the pelagic food web in Loch Ness

The natural abundance of stable isotopes (δ¹³C and δ¹³15N) was determined for components of the pelagic food web in Loch Ness, a deep oligotrophic lake in northern Scotland, and compared with values from the inflow rivers and the catchment vegetation. Phytoplankton δ¹³C was low compared to values reported from other lakes, possibly reflecting a high use of ¹³C-depleted carbon dioxide from respired organic matter before further isotopic fractionation during photosynthesis. Phytoplankton δ¹³C was appreciably lower than that of dissolved and particulate organic matter (DOM and POM) in the loch. The DOM and POM were evidently overwhelmingly of allochthonous origin and ultimately derived from terrestrial plant detritus. The distinctive δ¹³C values for phytoplankton and detritus in the loch allowed the use of food sources by grazing crustacean zooplankton to be assessed, and the contributions of phytoplankton carbon and detrital carbon to zooplankton total body carbon appeared to be about equal. Comparison of δ¹³C and δ¹⁵N values for zooplankton and fish allowed assessment of trophic structure in the loch. The very high dependence of the pelagic food web in Loch Ness on allochthonous organic matter inputs from the catchment may be exceptional in a large lake, but has important implications for our understanding of lake ecosystem processes as well as for lake management.     

Organic phosphorus in soil water under a European beech (Fagus sylvatica L.) stand in northeastern Bavaria, Germany: seasonal variability and changes with soil depth

Organically bound phosphorus (P) is a mobile form of phosphorus in many soils and thus its dynamics relevant for the leaching and cycling of this element. Despite its importance, little is known about the chemical composition of dissolved organic P. We studied the concentrations, fluxes, and chemical composition of organic P in forest floor leachates and soil solutions in a Rendzic Leptosol under a 90-year-old European beech (Fagus sylvatica L.) forest over a 27-month period (1997–1999). The chemical composition of organic P was analysed using XAD-8 fractionation and ³¹P-nuclear magnetic resonance (NMR) spectroscopy. Organic P was the dominant P form in forest floor leachates as well as in porewaters of the mineral soil. The largest concentrations of organic P were observed during summer and peaked (330–400 g dissolved organic P l^–1) after rain storms following short dry periods, concurrently with the concentrations of organic carbon (OC). Because of high rainfall, fluxes of organic P (and C) were greatest in autumn although concentrations of organic C and P were lower than in summer. In forest floor leachates, the hydrophilic fraction of dissolved organic matter contained 83 ± 13% of the bulk organic P. In soil solutions from 90 cm depth, organic P was almost exclusively in the hydrophilic fraction. Because of the low retention of the hydrophilic fraction of dissolved organic matter in the mineral soils, concentrations of organic P in soil water remained almost constant with depth. Consequently, organic P contributed > 95% of the total P leached into deeper subsoils. The overall retention of organic P in the weakly developed mineral soils was little and so the average annual fluxes of organic P in subsoils at 90 cm depth (38 mg m^–2) comprised 67% of those from the forest floors (57 mg m^–2) during the study period. Hence, organic P proved to be mobile in the studied soil. ³¹P-NMR spectroscopy confirmed the dominance of organic P species in soil water. Signals due to inorganic P occurred only in spectra of samples collected in winter and spring months. Spectra of samples from summer and autumn revealed traces of condensed phosphates. Due to low P contents, identification of organic P species in samples from winter and spring was not always possible. In summer and autumn, monoester and diester phosphates were the dominant organic species and varied little in their relative distributions. The distribution of organic species changed little from forest floor leachates to the subsoil solutions indicating that the composition of P-containing compounds was not influenced by sorptive interactions or biological transformation.     

杉木和米槠凋落叶DOM对土壤碳矿化的影响

DOM(Dissolved organic matter)是土壤微生物呼吸的重要底物,凋落物淋溶的DOM对土壤碳矿化具有重要影响。选择中亚热带地区具有代表性的杉木(Cunninghamia lanceolata)和米槠(Castanopsis carlesii)凋落叶作为研究对象,通过两个月的短期室内培养,把不同凋落叶浸提出的DOM添加到培养瓶中,定期测定土壤碳矿化速率,计算土壤碳累积矿化量,探讨两种等浓度等量DOM添加对土壤碳矿化的影响,并分析DOM化学性质在土壤碳矿化过程中的重要性。结果表明:米槠凋落叶浸提得到的DOC(Dissolved organic carbon)和DON(Dissolved organic nitrogen)浓度均显著高于杉木凋落叶的(P0.05),而杉木凋落叶浸提得到的DOM的UV吸收值(SUVA_(254))和HIX(Humification index)均显著低于米槠凋落叶的(P0.01)。添加等浓度等量杉木和米槠凋落叶DOM到土壤中均显著增加了土壤碳矿化速率,在第1天内分别比对照高198%和168%,3d后下降到61.8%和44.1%,14d后基本处于平稳状态,表明外源有机物添加对土壤碳矿化的前期影响较大。培养过程中,添加杉木和米槠凋落叶DOM的土壤碳矿化累积量均能采用双因素指数模型进行拟合(r~2=0.99),但添加两者凋落叶DOM后土壤碳矿化累积量没有显著差异。     

Tropical tree species composition affects the oxidation of dissolved organic matter from litter

Plant species effects on soil nutrient availability are relatively well documented, but the effects of species differences in litter chemistry on soil carbon cycling are less well understood, especially in the species-rich tropics. In many wet tropical forest ecosystems, leaching of dissolved organic matter (DOM) from the litter layer accounts for a significant proportion of litter mass loss during decomposition. Here we investigated how tree species differences in soluble dissolved organic C (DOC) and nutrients affected soil CO₂ fluxes in laboratory incubations. We leached DOM from freshly fallen litter of six canopy tree species collected from a tropical rain forest in Costa Rica and measured C-mineralization. We found significant differences in litter solubility and nutrient availability. Following DOM additions to soil, rates of heterotrophic respiration varied by as much as an order of magnitude between species, and overall differences in total soil CO₂ efflux varied by more than four-fold. Variation in the carbon: phosphorus ratio accounted for 51% of the variation in total CO₂ flux between species. These results suggest that tropical tree species composition may influence soil C storage and mineralization via inter-specific variation in plant litter chemistry.     

瓦屋山扁刺栲-中华木荷常绿阔叶次生林土壤有机碳组分特征

次生林在全球碳循环中占有重要地位,为了研究中国中亚热带次生林土壤有机碳组分特征,以四川瓦屋山中山段扁刺栲-中华木荷常绿阔叶次生林为对象,通过挖取土壤剖面分层(0—10、10—40、40—70 cm和70—100 cm)取样方式,研究土壤各有机碳组分特征。结果表明:土壤有机碳、微生物生物量碳、可浸提溶解性有机碳和易氧化碳含量均随土层深度增加而减小,0—10 cm土层有机碳含量为121.89 g/kg,高于已报道的亚热带其他常绿阔叶林和四川各类森林;0—10 cm层微生物生物量碳含量为1931.82 mg/kg,可浸提溶解性有机碳含量为697.42 mg/kg,易氧化碳含量为20.98 g/kg,高于已报道的许多相似天然林和人工林活性碳含量。土壤有机碳储量为154.87 t/hm2,在四川省各类森林中处于中等水平。研究表明瓦屋山扁刺栲-中华木荷常绿阔叶次生林活性碳含量较大,微生物活动和养分流动较为活跃,凋落物层转化为土壤碳的潜力较大,这类生态系统可能会在区域碳循环过程中扮演更为重要的角色。     

Dissolved organic sulphur in soil water under <Emphasis Type="Italic">Pinus sylvestris</Emphasis> L. and <Emphasis Type="Italic">Fagus sylvatica</Emphasis> L. stands in northeastern Bavaria,Germany variations with seasons and soil depth

Organically bound species have been identified as prominent and mobile forms of nitrogen and phosphorus in soils. Since a large portion of sulphur (S) in soil is bonded to carbon (C) also dissolved organic S likely is a significant constituent in soil water. To investigate the role of dissolved organic forms in leaching and cycling of S in forest soils, we examined concentrations, fluxes, and chemical composition of organic S in forest floor leachates and in soil solutions of Rendzic Leptosols under 90-year-old European beech (Fagus sylvatica L.) and Haplic Arenosols under 160-year-old Scots pine (Pinus sylvestris L.) for 27 months. These soils are low in adsorbed SO₄^2- and receive little atmospheric S depositions at present. The chemical composition of organic S was estimated by fractionation with XAD-8 and wet-chemical characterisation (HI reduction) of binding forms. Although not as prominent as the organic forms of other nutrient elements, organic S proved to be an important contributor to S dissolved in forest floor leachates and in mineral soil solutions. Dissolved organic matter contained on average 29% of total S in forest floor leachates at the pine site and 34% at the beech site. The largest portion of organic S occurred in the subsoil solutions under beech in summer and autumn (up to 53%). Mean concentrations of organic S peaked (up to 1.1 mg l^-1) in summer after rainstorms that followed dry periods. Fluxes with forest floor leachates and at 90 cm soil depth were largest in autumn because of huge amounts of rainfall. Organic S contributed significantly to the fluxes of S in the subsoils under beech comprising on average 39% of total dissolved S at 90 cm depth. Organic S produced in the forest floor layers was mainly in the hydrophilic fraction of dissolved organic matter (62 ± 6% at the pine site, 85 ± 4% at the beech site). The major binding form of organic S in the hydrophobic fraction was C-bonded S while in the hydrophilic fraction ester sulphate S, possibly associated with carbohydrates, was more prominent. Since the hydrophobic fraction increased in summer and autumn, C-bonded S was of greater importance during that time of the year than in winter and spring. With depth, concentrations and composition of organic S (and also of C) hardly changed at the pine site because of little retention of dissolved organic matter, presumably because of the small sorption capability of that soil. At the beech where organic C showed a marked decrease with depth, only a slight decrease in organic S, exclusively from the hydrophobic fraction, was found indicating that organic S was mobile compared with organic C. This was probably due to the concentration of S in the hydrophilic fraction of dissolved organic matter. Because of being concentrated in the mobile hydrophilic fraction, ester sulphate S was more mobile in the soil under beech than C-bonded S.     

Transport of Carbon and Nitrogen Between Litter and Soil Organic Matter in a Northern Hardwood Forest

We used sugar maple litter double-labeled with ¹³C and ¹⁵N to quantify fluxes of carbon (C) and nitrogen (N) between litter and soil in a northern hardwood forest and the retention of litter C and N in soil. Two cohorts of litter were compared, one in which the label was preferentially incorporated into non-structural tissue and the other structural tissue. Loss of ¹³C from this litter generally followed dry mass and total C loss whereas loss of ¹⁵N (20–30% in 1 year) was accompanied by large increases of total N content of this decaying litter (26–32%). Enrichment of ¹³C and ¹⁵N was detected in soil down to 10–15 cm depth. After 6 months of decay (November–May) 36–43% of the ¹³C released from the litter was recovered in the soil, with no differences between the structural and non-structural labeled litter. By October the percentage recovery of litter ¹³C in soil was much lower (16%). The C released from litter and remaining in soil organic matter (SOM) after 1 year represented over 30 g C m⁻² y⁻¹ of SOM accumulation. Recovery of litter ¹⁵N in soil was much higher than for C (over 90%) and in May ¹⁵N was mostly in organic horizons whereas by October it was mostly in 0–10 cm mineral soil. A small proportion of this N was recovered as inorganic N (2–6%). Recovery of ¹⁵N in microbial biomass was higher in May (13–15%) than in October (about 5%). The C:N ratio of the SOM and microbial biomass derived from the labeled litter was much higher for the structural than the non-structural litter and for the forest floor than mineral SOM, illustrating the interactive role of substrates and microbial activity in regulating the C:N stoichiometry of forest SOM formation. These results for a forest ecosystem long exposed to chronically high atmospheric N deposition (ca. 10 kg N ha⁻¹ y⁻¹) suggest possible mechanisms of N retention in soil: increased organic N leaching from fresh litter and reduced fungal transport of N from soil to decaying litter may promote N stabilization in mineral SOM even at a relatively low C:N ratio.     

Chemistry and Dynamics of Dissolved Organic Matter in a Temperate Coniferous Forest on Andic Soils: Effects of Litter Quality

Dissolved organic matter (DOM) plays an important role in transporting carbon and nitrogen from forest floor to mineral soils in temperate forest ecosystems. Thus, the retention of DOM via sorption or microbial assimilation is one of the critical steps for soil organic matter formation in mineral soils. The chemical properties of DOM are assumed to control these processes, yet we lack fundamental information that links litter quality, DOM chemistry, and DOM retention. Here, we studied whether differences in litter quality affect solution chemistry and whether changes in litter inputs affect DOM quality and removal in the field. The effects of litter quality on solution chemistry were evaluated using chemical fractionation methods for laboratory extracts and for soil water collected from a temperate coniferous forest where litter inputs had been altered. In a laboratory extraction, litter type (needle, wood, root) and the degree of decomposition strongly influenced solution chemistry. Root litter produced more than 10 times more water-extractable dissolved organic N (DON) than any other litter type, suggesting that root litter may be most responsible for DON production in this forest ecosystem. The chemical composition of the O-horizon leachate was similar under all field treatments (doubled needle, doubled wood, and normal litter inputs). O-horizon leachate most resembled laboratory extracts of well-decomposed litter (that is, a high proportion of hydrophobic acids), in spite of the significant amount of litter C added to the forest floor and a tendency toward higher mean DOM under doubled-Litter treatments. A lag in DOM production from added litter or microbial modification might have obscured chemical differences in DOM under the different treatments. Net DOM removal in this forest soil was strong; DOM concentration in the water deep in the mineral soil was always low regardless of concentrations in water that entered the mineral soil and of litter input manipulation. High net removal of DOM from O-horizon leachate, in spite of extremely low initial hydrophilic neutral content (labile DOM), coupled with the lack of influence by season or soil depth, suggests that DOM retention in the soil was mostly by abiotic sorption.     

Earthworms, litter and soil carbon in a northern hardwood forest

The important role of soil carbon (C) in the global C cycle has stimulated interest in better understanding the mechanisms regulating soil C storage and its stabilization. Exotic earthworm invasion of northern forest soils in North America can affect soil C pools, and we examined their effects on these mechanisms by adding ¹³C labeled leaf litter to adjacent northern hardwood forests with and without earthworms. Two types of labeled litter were produced, one with the ¹³C more concentrated in structural (S) components and the other in non-structural (NS) components, to evaluate the role of biochemical differences in soil C stabilization. Earthworm invasions have reduced soil C storage in the upper 20 cm of the soil profile by 37 %, mostly by eliminating surface organic horizons. Despite rapid mixing of litter into mineral soil and its incorporation into aggregates, mineral soil C has not increased in the presence of earthworms. Incorporation of litter C into soil and microbial biomass was not affected by biochemical differences between S versus NS labeled litter although NS litter C was assimilated more readily into earthworm biomass and S litter C into fungal hyphae. Apparently, the net effect of earthworm mixing of litter and forest floor C into mineral soil, plus stabilization of that C in aggregates, is counterbalanced by earthworm bioturbation and possible priming effects. Our results support recent arguments that biochemical recalcitrance is not a major contributor to the stabilization of soil C.     

Preferential use of root litter compared to leaf litter by beech seedlings and soil microorganisms

Background and aims

Litter decomposition is regulated by e.g. substrate quality and environmental factors, particularly water availability. The partitioning of nutrients released from litter between vegetation and soil microorganisms may, therefore, be affected by changing climate. This study aimed to elucidate the impact of litter type and drought on the fate of litter-derived N in beech seedlings and soil microbes.

Methods

We quantified ¹⁵N recovery rates in plant and soil N pools by adding ¹⁵N-labelled leaf and/or root litter under controlled conditions.

Results

Root litter was favoured over leaf litter for N acquisition by beech seedlings and soil microorganisms. Drought reduced ¹⁵N recovery from litter in seedlings thereby affecting root N nutrition. ¹⁵N accumulated in seedlings in different sinks depending on litter type.

Conclusions

Root turnover appears to influence (a) N availability in the soil for plants and soil microbes and (b) N acquisition and retention despite a presumably extremely dynamic turnover of microbial biomass. Compared to soil microorganisms, beech seedlings represent a very minor short-term N sink, despite a potentially high N residence time. Furthermore, soil microbes constitute a significant N pool that can be released in the long term and, thus, may become available for N nutrition of plants.