Impact of long-term N fertilization on the structural composition of spruce litter and mor humus

The aims of this study were to determine the degree of lignin degradation and to investigate changes in the chemical composition of the organic matter in the forest floor in an N fertilized Norway spruce forest soil. Needle litter and mor humus were collected from the field experiment at Skogaby in southern Sweden (56°33′N; 13°13′E). The spruce stand had been fertilized for 11 years with 100 kg N ha⁻¹ yr⁻¹ as (NH₄)₂SO₄. The degree of lignin degradation was determined with alkaline CuO oxidation followed by HPLC analysis. The chemical composition of the organic matter was characterized by CPMAS ¹³C NMR. Tannin was specifically analyzed using dipolar dephasing CPMAS ¹³C NMR and the N distribution was studied by CPMAS ¹⁵N NMR.The C-to-N ratios in the fertilized Oi and Oe layers were significantly lower than in the unfertilized layers (24 compared to 34 and 23 compared to 27, respectively). Neither the sum of the CuO oxidation products (Vanillyls+Syringyls+Cinnamyls expressed as VSC) nor the acid-to-aldehyde ratio ((Ac/Al)_V) showed any significant treatment effects. The content of aromatic C (including phenolic C) was significantly lower in the unfertilized than in the fertilized Oi layer (18 versus 21%). In the unfertilized soil, VSC was positively correlated (r=+0.63, p<0.05) with the C-to-N ratio, whereas the phenolic C content was negatively correlated (r=−0.61, p<0.05). The tannin index showed a tendency of increasing from Oi to Oe layers in both treatments. Most of the organic N was found as amide-N, whereas no heterocyclic N was detected. We have not been able to show any major C structural changes due to N fertilization. We suggest that the significantly higher content of aromatic and phenolic C in the fertilized Oi layer is due to an initial stimulation of the microbial community.     

Immobilization of avian excreta-derived nutrients and reduced lignin decomposition in needle and twig litter in a temperate coniferous forest

The possible effects of excreta of the Great Cormorant Phalacrocorax carbo on decomposition processes and dynamics of nutrients (N, P, Ca, K, Mg) and organic chemical components (lignin, total carbohydrates) were investigated in a temperate evergreen coniferous forest near Lake Biwa in central Japan. Two-year decomposition processes of needles and twigs of Chamaecyparis obtusa were examined at two sites, control site never colonized by the cormorants (site C) and colonizing site (site 2). Mass loss was faster in needles than in twigs. Mass loss of these litter types was faster at site C than at site 2, which was ascribed to the decreased mass loss rate of acid-insoluble ‘lignin’ at site 2. Net immobilization of N, P, and Ca occurred in needles and twigs at site 2; whereas at site C, mass of these elements decreased without immobilization during decomposition. Duration of immobilization phase of these nutrients at site 2 was estimated to be 1.6 to 2.5 years in needles and 19.6 to 23.5 years in twigs. Immobilization potential (maximum amount of exogenous nutrient immobilized per gram initial material) was similar between needles and twigs for N and Ca but was about 10 times higher in twigs than in needles for P. δ¹³C in needles was relatively constant during the first year and then increased during the second year, whereas δ¹³C in twigs was variable during decomposition. Acid-insoluble fraction was depleted in ¹³C compared to whole needles (1.6-2.1‰) and twigs (2.0-2.5‰). δ¹⁵N of needles and twigs and their acid-insoluble fractions approached to δ¹⁵N of excreta during decomposition at site 2. This result demonstrated the immobilization of excreta-derived N into litter due to the formation of acid-insoluble lignin-like substances complexed with excreta-derived N. No immobilization occurred in K and Mg and their mass decreased during decomposition at both sites. Based on these results of nutrient immobilization during decomposition and on the data of litter fall and excreta amount at site 2, we tentatively calculated stand-level immobilization potential of litter fall and its contribution to total amount of N and P deposited as excreta. Thus, the potential maximum amount immobilized into litter fall (needles and twigs) was estimated to account for 5-7% of total excreta-derived N and P.     

Ryegrass straw component decomposition during mesophilic and thermophilic incubations

The decomposition of perennial ryegrass straw was examined under mesophilic and thermophilic temperatures. Thermophilic conditions were used to define the composting process. The change in lipids, sugars, soluble polysaccharides, cellulose, and lignin was determined during a 45-day incubation. C, H, O, and N steadily decreased in both temperature treatments. The lignin content, as measured by the Klason or 72% H₂SO₄ method, decreased by 10% under mesophilic and 29% under thermophilic conditions. The Klason lignin C loss was 25 and 39% under mesophilic and thermophilic incubations, respectively. The changes in element (C, N, H, and O) ratios indicated that 94% of the lignin fraction was altered during both low- and high-temperature incubations. The changes in the lignin-like fraction as shown by elemental ratios were more extensive than those indicated by the Klason method, showing that this lignin determination has limited value in describing plant residue decomposition. The decomposition of the straw components and the concomitant degradation of the lignin fraction represent an important decomposition process that facilitates the composting of ryegrass straw with a high C:N ratio.     

CPMAS 13C NMR and IR spectra of spruce and pine litter and of the Klason lignin fraction at different stages of decomposition

Fresh and decomposed spruce and pine litter and the Klason lignin fraction of spruce needles at different stages of decomposition were studied by CPMAS ¹³C NMR and IR spectroscopy as well as by chemical methods. It was shown that decomposition of needles is accompanied by an increase in aliphatic substances and carboxyl group content; the amount of polysaccharides is reduced. It is assumed that stable aliphatic compounds like cutin and lipids of microbial origin will accumulate during litter decomposition and humification. Aromaticity is low and does not alter drastically. The NMR spectra of the Klason lignin fraction show pronounced peaks at 30, 55, 115, 130, 150 and 175 ppm. Obviously, this fraction contains appreciable amounts of aliphatic and carboxyl carbon besides the typical aromatic units of lignin. During decomposition aromaticity decreases whereas the relative amounts of aliphatic substances and carboxyl groups increase. This is probably due to splitting of aromatic ring structures and side chains. The findings agree with the results from chemical analyses.     

Saprotrophic fungi transform organic phosphorus from spruce needle litter

Fungal decomposition of and phosphorus transformation from spruce litter needles (Picea abies) were simulated in systems containing litter needles inoculated with individual saprotrophic fungal strains and their mixtures. Fungal strains of Setulipes androsaceus (L.) Antonín, Chalara longipes (Preus) Cooke, Ceuthospora pinastri (Fr.) Höhn., Mollisia minutella (Sacc.) Rehm, Scleroconidioma sphagnicola Tsuneda, Currah & Thormann and an unknown strain NK11 were used as representatives of autochthonous mycoflora. Systems were incubated for 5.5 months in laboratory conditions. Fungal colonization in systems and competition among strains were assessed using the reisolation of fungi from individual needles. After incubation, needles were extracted with NaOH and extracts were analysed using ³¹P nuclear magnetic resonance spectroscopy (NMR). Needle decomposition was determined based on the decrease in C:N ratio. Systems inoculated with the basidiomycete S. androsaceus revealed substantial decrease in C:N ratio (from 25.8 to 11.3) while the effect of ascomycetes on the C:N ratio was negligible. We suppose that tested strains of saprotrophic ascomycetes did not participate substantially in litter decomposition, but were directly involved in phosphorus transformation and together with S. androsaceus could transform orthophosphate monoesters and diesters from spruce litter needles into diphosphates, polyphosphates and phosphonates. These transformations seem to be typical for saprotrophic fungi involved in litter needle decomposition, although the proportion of individual phosphorus forms differed among studied fungal strains. Phosphonate presence in needles after fungal inoculation is of special interest because no previous investigation recorded phosphonate synthesis and accumulation by fungi. Our results confirmed that the ³¹P NMR spectroscopy is an excellent instrumental method for studying transformations of soil organic phosphorus during plant litter decomposition. We suggest that polyphosphate production by S. androsaceus may contribute to the phosphorus cycle in forest ecosystems because this fungus is a frequent litter colonizer that substantially participates in decomposition.     

Effects of nitrogen addition and litter properties on litter decomposition and enzyme activities of individual fungi

Litter decomposition is an important process of C and N cycling in the soil. Variation in the response of litter decomposition to nitrogen (N) addition (positive, negative or neutral) has been observed in many field studies. However, mechanism about variability in individual fungal species response to N addition has not yet been well demonstrated in the literature. Therefore, the objective of this study was to investigate the effects of N addition and litter chemistry properties on litter decomposition and enzyme activities of individual fungi. Three fungal species (Penicillium, Aspergillus, and Trichoderma) were isolated from a subtropical mixed forest soil. An incubation experiment was conducted using the individual fungi with two types of litter (leaf of Pinus massoniana and needle of Cryptocarya chinensis) and different N addition levels (0, 50 and 100 for N-deficient treatments, and 500 and 1000 μg N for N-excessive treatments). Cumulative CO₂-C, enzyme activities, and lignin and cellulose loss were measured during the incubation period of 60 days. Litter decomposition and enzyme activities significantly varied with the fungal species, while the N addition and litter types greatly affected fungal enzyme activities. The N treatments significantly increased lignin-rich needle decomposition by lignocellulose decomposers (Penicillium and Aspergillus) but did not affect their leaf decomposition. On the contrary, The N treatments stimulated leaf decomposition by cellulolytic species (Trichoderma) but did not affect its needle decomposition. Correlation analysis showed that lignin in the litter was the key component to affect litter decomposition. Activities of N-acetyl-β-glucosaminidase and phenol oxidase were both positively correlated to litter decomposition. The fungi (Penicillium and Aspergillus) with higher production of N-acetyl-β-glucosaminidase showed higher litter decomposition ability. The low N addition levels stimulated Penicillium and Aspergillus litter decomposition, but they still required more N source (e.g., litter N source) to support decomposition. Depressed fungal litter N uptake (lower N-acetyl-β-glucosaminidase activities) only occurred at the highest N addition level. Litter decomposition of Trichoderma depended more on external N and its litter decomposition capability was the lowest among the three species.     

Processes controlling the production of aromatic water-soluble organic matter during litter decomposition

Dissolved organic matter (DOM) plays a fundamental role for many soil processes. For instance, production, transport, and retention of DOM control properties and long-term storage of organic matter in mineral soils. Production of water-soluble compounds during the decomposition of plant litter is a major process providing DOM in soils. Herein, we examine processes causing the commonly observed increase in contribution of aromatic compounds to WSOM during litter decomposition, and unravel the relationship between lignin degradation and the production of aromatic WSOM. We analysed amounts and composition of water-soluble organic matter (WSOM) produced during 27 months of decomposition of leaves and needles (ash, beech, maple, spruce, pine). The contribution of aromatic compounds to WSOM, as indicated by the specific UV absorbance of WSOM, remained constant or increased during decomposition. However, the contribution of lignin-derived compounds to the total phenolic products of ¹³C-labelled tetramethylammonium hydroxide (¹³C-TMAH) thermochemolysis increased strongly (by >114%) within 27 months of decomposition. Simultaneous changes in contents of lignin phenols in solid litter residues (cupric oxide method as well as ¹³C-TMAH thermochemolysis) were comparably small (−39% to +21% within 27 months). This suggests that the increasing contribution of lignin-derived compounds to WSOM during decomposition does not reflect compositional changes of solid litter residues, but rather the course of decomposition processes. In the light of recently published findings, these processes include: (i) progressive oxidative alteration of lignin that results in increasing solubility of lignin, (ii) preferential degradation of soluble, non-lignin compounds that limits their contribution to WSOM during later phases of decomposition.     

Microbial biomass and activity in soil and litter underPinus sylvestris, Picea abies andBetula pendula at originally similar field afforestation sites

Microbial biomass C and N, and activities related to C and N cycles, were compared in needle and leaf litter, and in the uppermost 10 cm of soil under the litter layer in Scots pine (Pinus sylvestris L.), Norway spruce (Picea abies L.) and silver birch (Betula pendula L.) stands, planted on originally similar field afforestation sites 23–24 years ago. The ground vegetation was differentiated under different tree species, consisting of grasses and herbs under birch and pine, and mosses or no vegetation with a thick layer of needles under spruce. The C:N ratio of the soils was 13–21 and the soil pH_{CaCl 2} 3.8–5.2. Both showed little variation under different tree species. Microbial biomass C and N, C mineralization, net ammonification, reduction) did not differ significantly in soil under different tree species either. Birch leaf litter had a higher pH_{CaCl 2} (5.9) than spruce and pine needle litter (pH 5.0 and 4.8, respectively). The C:N ratio of spruce needles was 30, and was considerably higher in pine needles (69) and birch leaves (54). Birch leaves tended to have the highest microbial biomass C and C mineralization. Spruce needles appeared to have the highest microbial biomass N and net formation of mineral N, whereas formation of mineral N in pine needles and birch leaves was negligible. Microbial biomass C and N were of the same order of magnitude in the soil and litter samples but C mineralization was tenfold higher in the litter samples.     

The initial lignin:nitrogen ratio of litter from above and below ground sources strongly and negatively influenced decay rates of slowly decomposing litter carbon pools

Understanding the interactions between the initial biochemical composition and subsequent decomposition of plant litter will improve our understanding of its influence on microbial substrate use to explain the flow of organic matter between soil carbon pools. We determined the effects of land use (cultivation/native woodland/native pasture), litter type (above and below ground) and their interaction on the initial biochemical composition (carbon, nitrogen, water soluble carbon, lignin, tannin and cellulose) and decomposition of litter. Litter decomposition was studied as the mineralization of C from litter by microbial respiration and was measured as CO₂–C production during 105 d of laboratory incubation with soil. A two-pool model was used to quantify C mineralization kinetics. For all litter types, the active C pool decay rate constants ranged from 0.072 d⁻¹ to 0.805 d⁻¹ which represented relatively short half-lives of between 1 and 10 days, implying that this pool contained compounds that were rapidly mineralized by microbes during the initial stages of incubation. Conversely, the decay rate constants for the slow C pool varied widely between litter types within and among land uses ranging from 0.002 d⁻¹ and 0.019 d⁻¹ representing half-lives of between 37 and 446 days. In all litter types, the initial lignin:N ratio strongly and negatively influenced the decay rate of the slow C pool which implied that the interaction between these two litter quality variables had important controls over the decomposition of the litter slow C pool. We interpret our results to suggest that where the flow of C from the active pool to the slow pool is largely driven by microbial activity in soil, the rate of transfer of C will be largely controlled by the quality of litter under different land-use systems and particularly the initial lignin:N ratio of the litter. Compared with native pastures and cultivation, above and below ground litter from native woodland was characterized by higher lignin:N ratio and more slowly decomposing slow C pools which implies that litter is likely to persist in soils, however based on the sandy nature of the soils in this study, it is likely to lack protection from microbial degradation in the long term.     

Analytical pyrolysis of a soil profile under Scots pine

The chemical properties of pine needle litter cause slow decomposition, which results in an accumulation of highly lignocellulosic material on the forest floor. Decomposition of organic matter is important for the nutrient turnover in pine forests on nutrient-poor soils. We studied the biodegradation of needles in an organic layer focusing on the various stages of lignin degradation by fungi. Samples were obtained from pine needle litter and a stratified organic layer over nutrient-poor sand under a 60-year-old Scots pine (Pinus sylvestris forest stand. Pyrolysis mass spectrometry (PyMS) and pyrolysis gas chromatography mass spectrometry (PyGCMS) were used to characterize the chemical composition of the needles and the soil. The pyrolysis data show that diterpenoid acids are a main component in fresh needles, but rapidly decrease in the organic layer of the soil, as a result of decomposition. The chemical composition of the soil profile is dominated by guaiacyl lignin and polysaccharides from needle litter. The hexose/pentose ratio increases with depth in the soil profile. The partial preservation of hexose polymers is the result of the preferential decomposition of pentose polymers by white-rot fungi, and points to the input of microbially synthesized polysaccharides. Indications for the degradation of guaiacyl lignin are also found in the soil profile. Oxidative reactions by soil fungi result in a shortening of the side chain of the guaiacyl lignin derivatives and an increase of carbonyl and carboxyl groups. These degradational patterns of lignin in the soil profile under Scots pine are similar to those observed in lignin model compounds and wood lignin degraded by fungi under controlled laboratory conditions.     

Elevated CO<Subscript>2</Subscript> concentration affected pine and oak litter chemistry and the respiration and microbial biomass of soils amended with these litters

Elevated atmospheric CO₂ concentration ([CO₂]) may change litter chemistry which affects litter decomposability. This study investigated respiration and microbial biomass of soils amended with litter of Pinus densiflora (a coniferous species; pine) and Quercus variabilis (a deciduous species; oak) that were grown under different atmospheric [CO₂] and thus had different chemistry. Elevated [CO₂] increased lignin/N through increased lignin concentration and decreased N concentration. The CO₂ emission from the soils amended with litter produced under the same [CO₂] regime was greater for oak than pine litter, confirming that broadleaf litter with lower lignin decomposes faster than needle leaf litter. Within each species, however, soils amended with high lignin/N litter grown under elevated [CO₂] emitted more CO₂ than those with low lignin/N litter grown under ambient [CO₂]. Such contrasting effects of lignin/N on inter- and intra-species variations in litter decomposition should be ascribed to the effects of other litter chemistry variables including nonstructural carbohydrate, calcium and manganese as well as inhibitory effect of N on lignin decomposition. The microbial biomass was also higher in the soils amended with high lignin/N litter than those with low lignin/N litter probably due to low substrate use efficiency of lignin by microbes. Our study suggests that elevated [CO₂] increases lignin/N for both species, but increased lignin/N does not always reduce soil respiration and microbial biomass. Further study investigating a variety of tree species is required for more comprehensive understanding of inter- and intra-species variations of litter decomposition under elevated [CO₂].     

Long-term nitrogen additions increased surface soil carbon concentration in a forest plantation despite elevated decomposition

Forests cover one-third of the Earth’s land surface and account for 30-40% of soil carbon (C). Despite numerous studies, questions still remain about the factors controlling forest soil C turnover. Present understanding of global C cycle is limited by considerable uncertainty over the potential response of soil C dynamics to rapid nitrogen (N) enrichment of ecosystems, mainly from fuel combustion and fertilizer application. Here, we present a 15-year-long field study and show an average increase of 14.6% in soil C concentration in the 0-5 cm mineral soil layer in N fertilized (defined as N+ hereafter) sub-plots of a second-rotation Pinus radiata plantation in New Zealand compared to control sub-plots. The results of ¹⁴C and lignin analyses of soil C indicate that N additions significantly accelerate decomposition of labile and recalcitrant soil C. Using an annual-time step model, we estimated the soil C turnover time. In the N+ sub-plots, soil C in the light (a density < 1.70 g cm⁻³) and heavy fractions had the mean residence times of 23 and 67 yr, respectively, which are lower than those in the control sub-plots (36 and 133 yr in the light and heavy fractions, respectively). The commonly used lignin oxidation indices (vanillic acid to vanillin and syringic acid to syringaldehyde ratios) were significantly greater in the N+ sub-plots than in the control sub-plots, suggesting increased lignin decomposition due to fertilization. The estimation of C inputs to forest floor and δ¹³C analysis of soil C fractions indicate that the observed buildup of surface soil C concentrations in the N+ sub-plots can be attributed to increased inputs of C mass from forest debris. We conclude that long-term N additions in productive forests may increase C storage in both living tree biomass and soils despite elevated decomposition of soil organic matter.     

Microbial biomass and activity in soil and litter underPinus sylvestris, Picea abies andBetula pendula at originally similar field afforestation sites

Microbial biomass C and N, and activities related to C and N cycles, were compared in needle and leaf litter, and in the uppermost 10 cm of soil under the litter layer in Scots pine (Pinus sylvestris L.), Norway spruce (Picea abies L.) and silver birch (Betula pendula L.) stands, planted on originally similar field afforestation sites 23–24 years ago. The ground vegetation was differentiated under different tree species, consisting of grasses and herbs under birch and pine, and mosses or no vegetation with a thick layer of needles under spruce. The C:N ratio of the soils was 13–21 and the soil pH_{CaCl 2} 3.8–5.2. Both showed little variation under different tree species. Microbial biomass C and N, C mineralization, net ammonification, reduction) did not differ significantly in soil under different tree species either. Birch leaf litter had a higher pH_{CaCl 2} (5.9) than spruce and pine needle litter (pH 5.0 and 4.8, respectively). The C:N ratio of spruce needles was 30, and was considerably higher in pine needles (69) and birch leaves (54). Birch leaves tended to have the highest microbial biomass C and C mineralization. Spruce needles appeared to have the highest microbial biomass N and net formation of mineral N, whereas formation of mineral N in pine needles and birch leaves was negligible. Microbial biomass C and N were of the same order of magnitude in the soil and litter samples but C mineralization was tenfold higher in the litter samples.     

Litter- and ecosystem-driven decomposition under elevated CO2 and enhanced N deposition in a Sphagnum peatland

Peatlands represent massive global C pools and sinks. Carbon accumulation depends on the ratio between net primary production and decomposition, both of which can change under projected increases of atmospheric CO₂ and N deposition. The decomposition of litter is influenced by 1) the quality of the litter, and 2) the microenvironmental conditions in which the litter decomposes. This study aims at experimentally testing the effects of these two drivers in the context of global change. We studied the in situ litter decomposition from three common peatland species (Eriophorum vaginatum, Polytrichum strictum and Sphagnum fallax) collected after one year of litter production under pre-treatment conditions (elevated CO₂: 560 ppm or enhanced N: 3 g m⁻² y⁻¹ NH₄NO₃) and decomposed the following year under treatment conditions (same as pre-treatment). By considering the cross-effects between pre-treatments and treatments, we distinguished between the effects on mass loss of 1) the pre-treatment-induced litter quality and 2) the treatment conditions under which the litters were decomposing. The combination between CO₂ pre-treatment and CO₂ treatment reduced Polytrichum decomposition by −24% and this can be explained by litter quality-driven decomposition changes brought by the pre-treatment. CO₂ pre-treatment reduced Eriophorum litter quality, although this was not sufficient to predict decomposition. The N addition pre-treatment reduced the decomposition of Eriophorum, due to enhanced lignin and soluble phenols concentrations in the initial litter, and reduced litter-driven losses of starch and enhanced litter-driven losses of soluble phenols. While decomposition indices based on initial litter quality provide a broad explanation of quantitative and qualitative decomposition, they can only be taken as first approximations. Indeed, the microbial ATP activity, the litter N loss and resulting litter quality, were strongly altered irrespective of the compounds' initial concentration and by means of processes that occurred independently of the initial litter-qualitative changes. The experimental design was valuable to assess litter- and ecosystem-driven decomposition pathways simultaneously or independently. The ability to separate these two drivers makes it possible to attest the presence of litter-qualitative changes even without any litter biochemical determinations, and shows the screening potential of this approach for future experiments dealing with multiple plant species.     

Leaf litter C and N cycling from a deciduous permanent crop

Our understanding of leaf litter carbon (C) and nitrogen (N) cycling and its effects on N management of deciduous permanent crops is limited. In a 30-day laboratory incubation, we compared soil respiration and changes in mineral N [ammonium (NH₄⁺-N) + nitrate (NO₃^–-N)], microbial biomass nitrogen (MBN), total organic carbon (TOC) and total non-extractable organic nitrogen (TON) between a control soil at ¹⁵N natural abundance (δ¹⁵N = 1.08‰) without leaf litter and a treatment with the same soil, but with almond (Prunus dulcis (Mill.) D.A. Webb) leaf litter that was also enriched in ¹⁵N (δ¹⁵N = 213‰). Furthermore, a two-end member isotope mixing model was used to identify the source of N in mineral N, MBN and TON pools as either soil or leaf litter. Over 30 d, control and treatment TOC pools decreased while the TON pool increased for the treatment and decreased for the control. Greater soil respiration and significantly lower (p < 0.05) mineral N from 3 to 15 d and significantly greater MBN from 10 to 30 d were observed for the treatment compared to the control. After 30 d, soil-sourced mineral N was significantly greater for the treatment compared to the control. Combined mineral N and MBN pools derived from leaf litter followed a positive linear trend (R² = 0.75) at a rate of 1.39 μg N g^?1 soil day^?1. These results suggest early-stage decomposition of leaf litter leads to N immobilization followed by greater N mineralization during later stages of decomposition. Direct observations of leaf litter C and N cycling assists with quantifying soil N retention and availability in orchard N budgets.     

Microbial community response to nitrogen deposition in northern forest ecosystems

The productivity of temperate forests is often limited by soil N availability, suggesting that elevated atmospheric N deposition could increase ecosystem C storage. However, the magnitude of this increase is dependent on rates of soil organic matter formation as well as rates of plant production. Nonetheless, we have a limited understanding of the potential for atmospheric N deposition to alter microbial activity in soil, and hence rates of soil organic matter formation. Because high levels of inorganic N suppress lignin oxidation by white rot basidiomycetes and generally enhance cellulose hydrolysis, we hypothesized that atmospheric N deposition would alter microbial decomposition in a manner that was consistent with changes in enzyme activity and shift decomposition from fungi to less efficient bacteria. To test our idea, we experimentally manipulated atmospheric N deposition (0, 30 and 80 kg NO₃⁻-N) in three northern temperate forests (black oak/white oak (BOWO), sugar maple/red oak (SMRO), and sugar maple/basswood (SMBW)). After one year, we measured the activity of ligninolytic and cellulolytic soil enzymes, and traced the fate of lignin and cellulose breakdown products (¹³C-vanillin, catechol and cellobiose).In the BOWO ecosystem, the highest level of N deposition tended to reduce phenol oxidase activity (131±13 versus 104±5 μmol h⁻¹ g⁻¹) and peroxidase activity (210±26 versus 190±21 μmol h⁻¹ g⁻¹) and it reduced ¹³C-vanillin and ¹³C-catechol degradation and the incorporation of ¹³C into fungal phospholipids (p<0.05). Conversely, in the SMRO and SMBW ecosystems, N deposition tended to increase phenol oxidase and peroxidase activities and increased vanillin and catechol degradation and the incorporation of isotope into fungal phospholipids (p<0.05). We observed no effect of experimental N deposition on the degradation of ¹³C-cellulose, although cellulase activity showed a small and marginally significant increase (p<0.10). The ecosystem-specific response of microbial activity and soil C cycling to experimental N addition indicates that accurate prediction of soil C storage requires a better understanding of the physiological response of microbial communities to atmospheric N deposition.     

Carbon and nitrogen turnover in response to warming and nitrogen addition during early stages of forest litter decomposition—an incubation experiment

Purpose

Little is known about the interactive effects of temperature, nitrogen (N) supply, litter quality, and decomposition time on the turnover of carbon (C) and N of forest litter. The objective of this study was to investigate the interactive effects of warming, N addition and tree species on the turnover of C and N during the early decomposition stage of litters in a temperate forest.

Materials and methods

A 12-week laboratory incubation experiment was carried out. The leaf litters including two types of broadleaf litters (Quercus mongolica and Tilia amurensis), a needle litter (Pinus koraiensis), and a mixed litter of them were collected from a broad-leaved Korean pine mixed forest ecosystem in northeastern China in September 2009. Nine treatments were conducted using three temperatures (15, 25, and 35 °C) combined with three doses of N addition (equal to 0, 75, and 150 kg?·?ha^?1?a^?1, respectively, as NH₄NO₃).

Results and discussion

After 12 weeks of incubation, the mass loss ranged between 12 and 35 %. The broadleaf litters had greater mass loss and cumulative CO₂–C emission than the needle litter. Temperature and N availability interacted to affect litter mass loss and decomposition rate. The dissolved organic carbon (DOC) and nitrogen (DON) concentrations in litter leachate varied widely with litter types. DOC increased significantly with increased temperature but decreased significantly with increased N availability. DON increased significantly with increased N availability but showed a higher level at the moderate decomposition temperature. The amounts of CO₂ and N₂O emission were significantly higher at 25 °C than those at 15 and 35 °C, and were significantly increased by the N addition.

Conclusions

The present study indicated relatively intricate temperature and N addition effects on C and N cycling during early stages of litter decomposition, implying that future increases in temperature and N deposition will directly affect C and N cycling in broad-leaved Korean pine mixed forest ecosystem, and may indirectly influence the ecosystem composition, productivity, and functioning in NE China. It is, therefore, important to understand the interactive effects of biotic and abiotic factors on litter decomposition in field conditions in order to assess and predict future ecosystem responses to environmental changes in NE China.     

Carbon fluxes in soil food webs of increasing complexity revealed by C labelling and C natural abundance

Soil food webs are mainly based on three primary carbon (C) sources: root exudates, litter, and recalcitrant soil organic matter (SOM). These C sources vary in their availability and accessibility to soil organisms, which could lead to different pathways in soil food webs. The presence of three C isotopes (¹²C, ¹³C and ¹⁴C) offers an unique opportunity to investigate all three C sources simultaneously. In a microcosm experiment we studied the effect of food web complexity on the utilization of the three carbon sources. We choose an incomplete three factorial design with (i) living plants, (ii) litter and (iii) food web complexity. The most complex food web consisted of autochthonous microorganisms, nematodes, collembola, predatory mites, endogeic and anecic earthworms. We traced C from all three sources in soil, in CO₂ efflux and in individual organism groups by using maize grown on soil developed under C₃ vegetation and application of ¹⁴C labelled ryegrass shoots as a litter layer. The presence of living plants had a much greater effect on C pathways than food web complexity. Litter decomposition, measured as ¹⁴CO₂ efflux, was decreased in the presence of living plants from 71% to 33%. However, living plants increased the incorporation of litter C into microbial biomass and arrested carbon in the litter layer and in the upper soil layer. The only significant effect of food web complexity was on the litter C distribution in the soil layers. In treatments with fungivorous microarthropods (Collembola) the incorporation of litter carbon into mineral soil was reduced. Root exudates as C source were passed through rhizosphere microorganisms to the predator level (at least to the third trophic level). We conclude that living plants strongly affected C flows, directly by being a source of additional C, and indirectly by modifying the existing C flows within the food web including CO₂ efflux from the soil and litter decomposition.     

C-NMR analysis of decomposing litter and fine roots in the semi-arid Mulga Lands of southern Queensland

Plant litter and fine roots are important carbon (C) inputs to soil and a direct source of CO₂ to the atmosphere. Solid-state carbon-13 nuclear magnetic resonance (¹³C-NMR) spectroscopy was used to investigate the nature of C changes during decomposition of plant litter and fine roots of mulga (Acacia aneura F. Muell. Ex. Benth.), wheat (Triticum aestivum L.), lucerne (Medicago sativa) and buffel grass (Cenchrus ciliaris) over an 18-month period. Alkyl C was closely associated with total N concentrations in all litter materials during decay and as alkyl C increased so did total N, indicating an increase in refractory biomacromolecules. Mulga phyllodes had the greatest alkyl C concentration of all litter and fine root materials, and also exhibited the NMR peaks assigned to tannins that may slow or hinder decomposition rates and nitrification. Mulga litter and fine roots decomposed slower than all other litter materials and the soil under mulga had the highest soil C concentration, indicating slower CO₂ release. The alkyl C-to-O-alkyl C ratio is generally used as an index of the extent of decomposition, but is not useful for the decay of woody components. Of all the NMR ratios studied that may indicate the extent of decomposition, the carbohydrate C-to-methoxyl C ratio proved to have the strongest and most consistent relationship with decay time, fraction of mass remaining and total C, even though increases in alkyl C were observed with decreases in carbohydrate C.     

Atmospheric nitrate deposition and the microbial degradation of cellobiose and vanillin in a northern hardwood forest

Human activity has increased the amount of N entering terrestrial ecosystems from atmospheric NO₃⁻ deposition. High levels of inorganic N are known to suppress the expression of phenol oxidase, an important lignin-degrading enzyme produced by white-rot fungi. We hypothesized that chronic NO₃⁻ additions would decrease the flow of C through the heterotrophic soil food web by inhibiting phenol oxidase and the depolymerization of lignocellulose. This would likely reduce the availability of C from lignocellulose for metabolism by the microbial community. We tested this hypothesis in a mature northern hardwood forest in northern Michigan, which has received experimental atmospheric N deposition (30 kg NO₃⁻-N ha⁻¹ y⁻¹) for nine years. In a laboratory study, we amended soils with ¹³C-labeled vanillin, a monophenolic product of lignin depolymerization, and ¹³C-labeled cellobiose, a disaccharide product of cellulose degradation. We then traced the flow of ¹³C through the microbial community and into soil organic carbon (SOC), dissolved organic carbon (DOC), and microbial respiration. We simultaneously measured the activity of enzymes responsible for lignin (phenol oxidase and peroxidase) and cellobiose (β-glucosidase) degradation. Nitrogen deposition reduced phenol oxidase activity by 83% and peroxidase activity by 74% when compared to control soils. In addition, soil C increased by 76%, whereas microbial biomass decreased by 68% in NO₃⁻ amended soils. ¹³C cellobiose in bacterial or fungal PLFAs was unaffected by NO₃⁻ deposition; however, the incorporation of ¹³C vanillin in fungal PLFAs extracted from NO₃⁻ amended soil was 82% higher than in the control treatment. The recovery of ¹³C vanillin and ¹³C cellobiose in SOC, DOC, microbial biomass, and respiration was not different between control and NO₃⁻ amended treatments. Chronic NO₃⁻ deposition has stemmed the flow of C through the heterotrophic soil food web by inhibiting the activity of ligninolytic enzymes, but it increased the assimilation of vanillin into fungal PLFAs.