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.     

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.     

Chemistry of soil organic matter as related to C : N in Norway spruce forest (Picea abies(L.) Karst.) floors and mineral soils

Due to high nitrogen deposition in central Europe, the C : N ratio of litter and the forest floor has narrowed in the past. This may cause changes in the chemical composition of the soil organic matter. Here we investigate the composition of organic matter in Oh and A horizons of 15 Norway spruce soils with a wide range of C : N ratios. Samples are analyzed with solid‐state ¹³C nuclear magnetic resonance (NMR) spectroscopy, along with chemolytic analyses of lignin, polysaccharides, and amino acid‐N. The data are investigated for functional relationships between C, N contents and C : N ratios by structural analysis. With increasing N content, the concentration of lignin decreases in the Oh horizons, but increases in the A horizons. A negative effect of N on lignin degradation is observed in the mineral soil, but not in the humus layer. In the A horizons non‐phenolic aromatic C compounds accumulate, especially at low N values. At high N levels, N is preferentially incorporated into the amino acid fraction and only to a smaller extent into the non‐hydrolyzable N fraction. High total N concentrations are associated with a higher relative contribution of organic matter of microbial origin.     

Coarse particulate organic matter is the primary source of mineral N in the topsoil of three beech forests

Forest soils contain a variable amount of organic N roughly repartitioned among particles of different size, microbial biomass and associated with mineral compounds. All pools are alimented by annual litter fall as main input of organic N to the forest floor. Litter N is further subject to mineralization/stabilization recognized as the crucial process for the turnover of litter N. Although it is well documented that different soil types have different soil N stocks, it is presently unknown how different soil types affect the turnover of recent litter N. Here, we compared the potential mineralization of the total soil organic N with that of recent litter-released N in three beech forests varying in their soil properties. Highly ¹⁵N-labelled beech litter was applied to stands located at Aubure, Ebrach, Collelongo, which differ in humus type, soil type and soil chemistry. After 4-5 years of litter decomposition, the upper 3 cm of the organo-mineral A horizon was sampled and the net N mineralization was measured over 112 days under controlled conditions. The origin of mineralized N (litter N versus soil organic N) was calculated using ¹⁵N labeling. In addition, soils were fractionated according to their particle size (>2000 μm, 200-2000 μm, 50-200 μm, <50 μm) and particulate organic matter (POM) was separated from the mineral fraction in size classes, except the <50 μm fraction. Between 41 and 69% of soil organic N was recovered as POM. Litter-released ¹⁵N was mainly to be found in the coarse POM fractions >200 μm. On a soil mass basis, N mineralization was two-fold higher at Aubure and Collelongo than at Ebrach, but, on a soil N basis, N mineralization was the lowest at Collelongo and the highest at Ebrach. On a soil N (or ¹⁵N) basis, mineralization of litter ¹⁵N was two to four-fold higher than mineralization of the average soil N. Furthermore, the δ¹⁵N of the mineral N produced was closer to that of POM than to that of the mineral-bound fraction (<50 μm). Highest rates of ¹⁵N mineralization happened in the soil with the lowest N content, and we found a negative relationship between accumulations of N in the upper A horizon and the mineralization of ¹⁵N from the litter. Our results show that mineral N is preferentially mineralized from POM in the upper organo-mineral soil irrespective of the soil chemistry and that the turnover rate of litter N is faster in soils with a low N content.     

Impacts of anthropogenic N additions on nitrogen mineralization from plant litter in exotic annual grasslands

Urban regions of southern California receive up to 45 kg N ha^-1 y^-1 from nitrogen (N) deposition. A field decomposition study was done using ¹⁵N-labelled litter of the widespread exotic annual grass Bromus diandrus to determine whether elevated soil N is strictly from N deposition or whether N mineralization rates from litter are also increased under N deposition. Tissue N and lignin concentrations, which are inversely related in field sites with high and low N deposition, determine the rate at which N moves from plant litter to soil and becomes available to plants. The effect of soil N on N movement from litter to soil was tested by placing litter on high and low N soil in a factorial experiment with two levels of litter N and two levels of soil N. The litter quality changes associated with N deposition resulted in faster rates of N cycling from litter to soil. Concentrations of litter-derived N in total N, NH₄⁺, NO₃⁻, microbial N and organic N were all higher from high N/low lignin litter than from low N/high lignin litter. Litter contributed more N to soil NH₄⁺ and microbial N in high N than low N soil. At the end of the study, N mineralized from high N litter on high N soil accounted for 46% of soil NH₄⁺ and 11% of soil NO₃⁻, compared to 35% of soil NH₄⁺ and 6% of soil NO₃⁻ from low N litter on low N soil. The study showed that in high N deposition areas, elevated inorganic soil N concentrations at the end of the summer N deposition season are a result of N mineralized from plant litter as well as from N deposition.     

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.     

Clear-cutting of a Norway spruce stand: implications for controls on the dynamics of dissolved organic matter in the forest floor

Clear‐cutting of forest provides a unique opportunity to study the response of dynamic controls on dissolved organic matter. We examined differences in concentrations, fluxes and properties of dissolved organic matter from a control and a clear‐cut stand to reveal controlling factors on its dynamics. We measured dissolved organic C and N concentrations and fluxes in the Oi, Oe and Oa horizons of a Norway spruce stand and an adjacent clear‐cutting over 3 years. Aromaticity and complexity of organic molecules were determined by UV and fluorescence spectroscopy, and we measured δ¹³C ratios over 1 year. Annual fluxes of dissolved organic C and N remained unchanged in the thin Oi horizon (～ 260 kg C ha^?1, ～ 8.5 kg N ha^?1), despite the large reduction in fresh organic matter inputs after clear‐cutting. We conclude that production of dissolved organic matter is not limited by lack of resource. Gross fluxes of dissolved organic C and N increased by about 60% in the Oe and 40% in the Oa horizon upon clear‐cutting. Increasing organic C and N concentrations and increasing water fluxes resulted in 380 kg C ha^?1 year^?1 and 10.5 kg N ha^?1 year^?1 entering the mineral soil of the clear‐cut plots. We found numerous indications that the greater microbial activity induced by an increased temperature of 1.5°C in the forest floor is the major factor controlling the enhanced production of dissolved organic matter. Increasing aromaticity and complexity of organic molecules and depletion of ¹³C pointed to an accelerated processing of more strongly decomposed parts of the forest floor resulting in increased release of lignin‐derived molecules after clear‐cutting. The largest net fluxes of dissolved organic C and N were in the Oi horizon, yet dissolved organic matter sampled in the Oa horizon did not originate mainly from the Oi horizon. Largest gross fluxes in the Oa horizon (control 282 kg C ha^?1) and increased aromaticity and complexity of the molecules with increasing depth suggested that dissolved organic matter was derived mainly from decomposition, transformation and leaching of more decomposed material of the forest floor. Our results imply that clear‐cutting releases additional dissolved organic matter which is sequestered in the mineral soil where it has greater resistance to microbial decay.     

Tree pruning mulch increases soil C and N in a shaded coffee agroecosystem in Hawaii

Agroforestry can increase the sequestration of carbon (C) in soils of tropical agroecosystems through increased litter and tree pruning inputs. Decomposition of these inputs is a key process in the formation of soil organic matter and in nutrient cycling. Our objectives were to study decay of tree pruning mulch and effects on soil C and N in a shaded coffee agroecosystem in Hawaii. Chipped tree pruning residues (mulch) were added to coffee plots shaded with the Leucaena hybrid KX2 over three years. We measured mulch decomposition and nitrogen loss over one year and changes in soil carbon and nitrogen (N) over two years. Mass loss of mulch was 80% over one year and followed first-order decay dynamics. There was significant loss from all major biochemical components. Net N loss from the mulch was positive throughout the entire period. The C:N and lignin:N ratios of the mulch declined significantly over the decomposition period. Mulch additions significantly increased soil C and N in the top 20 cm by 10.8 and 2.12 Mg ha⁻¹, respectively. In the no-mulch treatment, there was no significant change in soil C or N concentration, but a decline in soil bulk density led to a significant decline in total soil C. Leucaena mulch can provide an important source of organic C and N to coffee agroecosystems and can help sequester C lost as plant biomass during shade tree management.     

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.     

川西亚高山人工云杉林地有机物和养分库的退化与调控

研究了川西亚高山云杉人工林地有机物和养分库状况 ,结果表明 :该区云杉人工林有机物和养分库严重退化 ,表现为 ,其凋落物的分解速率和周转期均较次生阔叶林和原始云杉林慢 ,致使地表枯枝落叶干物质和各种养分贮量滞留于凋落物层而不能进入土壤 ,土壤中有机质、全N、全P和碱解N含量随人工云杉林龄的增加而大幅度下降。人工云杉林份组成单一 ,其凋落物分解慢 ,归还土壤凋落物和养分数量少 ,是川西亚高山云杉人工林地土壤有机物和养分库退化的重要原因 ,人为收集凋落物积肥和人工抚育清灌 ,不断带走植被中养分是土壤有机物和养分库不断耗竭的另一重要原因。建议对该区人工成熟林抚育间伐和营造针阔混交林 ,改善成熟林下微环境和改变林份组成 ,可在很大程度上防治云杉人工林土壤有机物和养分库的退化     

Soil organic matter quantity and quality in mountain soils of the Alay Range, Kyrgyzia, affected by land use change

 Changes in soil management practices influence the amount, quality and turnover of soil organic matter (SOM). Our objective was to study the effects of deforestation followed by pasture establishment on SOM quantity, quality and turnover in mountain soils of the Sui Checti valley in the Alay Range, Kyrgyzia. This objective was approached by analysis of total organic C (TOC), N, lignin-derived phenols, and neutral sugars in soil samples and primary particle-size soil fractions. Pasture installation led to a loss of about 30% TOC compared with the native Juniperus turkestanica forests. The pasture soils accumulated about 20% N, due to inputs via animal excrement. A change in land use from forest to pasture mainly affected the SOM bound to the silt fraction; there was more microbial decomposition in the pasture than in the forest silt fraction, as indicated by lower yields of lignin and carbohydrates, and also by a more advanced oxidative lignin side-chain oxidation and higher values of plant : microbial sugar ratios. The ratio of arabinose : xylose was indicative of the removal of carbohydrates when the original forest was replaced by pasture, and we conclude that this can be used as an indicator of deforestation. The accumulation of lignin and its low humification within the forest floor could be due to the extremely cold winter and dry summer climate. Received: 10 March 1999     

Forest floor development and biochemical properties in reconstructed boreal forest soils

Following resource extraction by surface mining in the oil sands region of northeastern Alberta, sites are reclaimed by reconstructing soils using a variety of salvaged organic and mineral materials, and planted to native tree species. This study assessed the influence of three distinct stand types (Populus tremuloides Michx., Pinus banksiana Lamb., and Picea glauca (Moench) Voss) on forest floor development (thickness, morphology, total carbon and nitrogen contents), soil organic matter composition, and associated soil microbial communities. Forest floor and top mineral soil (0–5 cm) samples were collected from 32 sites reclaimed 16–33 years ago. Soil organic matter composition was measured using ramped-cross-polarization ¹³C nuclear magnetic resonance, and microbial communities were characterized using phospholipid fatty acid analysis. Morphological characteristics indicated little mesofaunal or fungal activities within the forest floors. Stands dominated by P. tremuloides fostered more rapid forest floor development than the coniferous (P. banksiana and P. glauca) stands, and showed a significant increase in forest floor thickness with time since reclamation. Within the P. tremuloides stands, forest floor development was accompanied by temporal changes in soil organic matter composition that reflected inputs from the canopy. Soil microbial community composition differed among reclamation treatments of the reconstructed soils, specifically as a function of their subsoil mineral textures, when canopy cover was below 30%. Above 30%, significant differences became apparent among stand types. Taken together, our results document how canopy cover and stand type were both important factors for the reestablishment of plant–soil relationships at these sites. Furthermore, achieving a canopy cover of 30% emerged as a critical threshold point during soil reclamation.     

Lignin degradation during a laboratory incubation followed by 13C isotope analysis

Recent in situ ¹³C studies suggest that lignin is not stabilised in soil in its polymerised form. However, the fate of its transformation products remains unknown. The objective of the present research was to provide the first comprehensive picture of the fate of lignin-derived C across its transformations processes: (1) C remaining as undecomposed lignin molecules, (2) C in newly formed humic substances, i.e. no longer identifiable as lignin-polymer C, (3) C in microbial biomass, (4) C mineralised as CO₂, and (5) dissolved organic C. To achieve this objective, we designed an incubation experiment with ¹³C-labelled lignin where both elementary and molecular techniques were applied. Lignin was isolated from ¹³C labelled maize plants (¹³C-MMEL) and incubated in an agricultural soil for 44 weeks. Carbon mineralisation and stable isotope composition of the released CO₂ were monitored throughout the incubation. Microbial utilisation of ¹³C-MMEL was measured seven times during the experiment. The turnover rate of the lignin polymer was assessed by ¹³C analysis of CuO oxidation products of soil lignin molecules. After 44 incubation weeks, 6.0% of initial ¹³C-MMEL carbon was mineralised, 0.8% was contained in the microbial biomass, and 0.1% was contained in dissolved organic C form. The compound-specific ¹³C data suggest that the remaining 93% were overwhelmingly in the form of untransformed lignin polymer. However, limited transformation into other humic substances potentially occurred, but could not be quantified because the yield of the CuO oxidation method proved somewhat variable with incubation time. The initial bacterial growth yield efficiency for MMEL was 31% and rapidly decreased to plateau of 8%. A two-pool first-order kinetics model suggested that the vast majority (97%) of MMEL lignin had a turnover time of about 25 years, which is similar to field-estimated turnover times for soil-extractable lignin but much longer than estimated turnover times for fresh plant-residue lignin. We conclude that natural lignin structures isolated from plants are rather unreactive in soil, either due to the lack of easily available organic matter for co-metabolism or due to enhanced adsorption properties. The data also suggest that fairly undecomposed lignin structures are the main reservoir of lignin-derived C in soils.     

Post-harvest residue management effects on recalcitrant carbon pools and plant biomarkers within the soil heavy fraction in Pinus radiata plantations

Forest soils contain about 30% of terrestrial carbon (C) and so knowledge of the influence of forest management on stability of soil C pools is important for understanding the global C cycle. Here we present the changes of soil C pools in the 0-5 cm layer in two second-rotation Pinus radiata (D.Don) plantations which were subjected to three contrasting harvest residue management treatments in New Zealand. These treatments included whole-tree harvest plus forest floor removal (defined as forest floor removal hereafter), whole-tree, and stem-only harvest. Soil samples were collected 5, 10 and 15 years after tree planting at Kinleith Forest (on sandy loam soils) and 4, 12 and 20 years after tree planting at Woodhill Forest (on sandy soils). These soils were then physically divided into light (labile) and heavy (stable) pools based on density fractionation (1.70 g cm⁻³). At Woodhill, soil C mass in the heavy fraction was significantly greater in the whole-tree and stem-only harvest plots than the forest floor removal plots in all sampling years. At Kinleith, the soil C mass in the heavy fraction was also greater in the stem-only harvest plots than the forest floor removal plots at year 15. The larger stable soil C pools with increased residue return was supported by analyses of the chemical composition and plant biomarkers in the soil organic matter (SOM) heavy fractions using NMR and GC/MS. At Woodhill, alkyl C, cutin-, suberin- and lignin-derived C contents in the SOM heavy fraction were significantly greater in the whole-tree and stem-only harvest plots than in the forest floor removal plots in all sampling years. At Kinleith, alkyl C (year 15), cutin-derived C (year 5 and 15) and lignin-derived C (Year 5 and 10) contents in the SOM heavy fraction were significantly greater in stem-only harvest plots than in plots where the forest floor was removed. The analyses of plant C biomarkers and soil δ¹³C in the light and heavy fractions of SOM indicate that the increased stable soil C in the heavy fraction with increased residue return might be derived from a greater input of recalcitrant C in the residue substrate.     

Comparison of forest soil carbon dynamics at five sites along a latitudinal gradient

The aim of this study was to compare the turnover time of labile soil carbon (C), in relation to temperature and soil texture, in several forest ecosystems that are representative of large areas of North America. Carbon and nitrogen (N) stocks, and C:N ratios, were measured in the forest floor, mineral soil, and two mineral soil fractions (particulate and mineral-associated organic matter, POM and MOM, respectively) at five AmeriFlux sites along a latitudinal gradient in the eastern United States. Sampling at four sites was replicated over two consecutive years. With one exception, forest floor and mineral soil C stocks increased from warm, southern sites (with fine-textured soils) to cool, northern sites (with more coarse-textured soils). The exception was a northern site, with less than 10% silt-clay content, that had a soil organic C stock similar to the southern sites. A two-compartment model was used to calculate the turnover time of labile soil organic C (MRT_U) and the annual transfer of labile C to stable C (k₂) at each site. Moving from south to north, MRT_U increased from approximately 5 to 14 years. Carbon-13 enrichment factors (ε), that described the rate of change in δ¹³C through the soil profile, were associated with soil C turnover times. Consistent with its role in stabilization of soil organic C, silt-clay content was positively correlated (r = 0.91; P ≤ 0.001) with parameter k₂. Latitudinal differences in the storage and turnover of soil C were related to mean annual temperature (MAT, °C), but soil texture superseded temperature when there was too little silt and clay to stabilize labile soil C and protect it from decomposition. Each site had a relatively high proportion of labile soil C (nearly 50% to a depth of 20 cm). Depending on unknown temperature sensitivities, large labile pools of forest soil C are at risk of decomposition in a warming climate, and losses could be disproportionately higher from coarse textured forest soils.     

Plant lignin and nitrogen contents control carbon dioxide production and nitrogen mineralization in soils incubated with Bt and non-Bt corn residues

Bt (Bacillus thuringiensis) corn is reported to produce lignin-rich residues, compared to non-Bt (NBt) corn, suggesting it is more resistant to decomposition. As the Bt gene is expressed selectively in stem and leaf tissue, it could affect lignin distribution in corn, which naturally has greater lignin content in roots than in stems and leaves. Our objective was to evaluate the effects of corn plant components, the Bt gene and elevated-lignin inputs on decomposition. Roots, stems and leaves from Bt corn and NBt corn isolines enriched with ¹³C and ¹⁵N were finely ground and mixed separately with soil, then incubated at 20 °C for 36 weeks. The effect of elevated lignin on decomposition was tested by adding a commercial lignin source (indulin lignin) to half of the samples. In addition to weekly CO₂ analysis and regular measurement of N mineralization, the degree of lignin degradation was evaluated at 1 and 36 weeks from the acid to aldehyde ratio (Ad/Al) of vanillyl and syringyl lignin-derived phenols. The CO₂ production and N mineralization was lower in root-amended soils than stem- and leaf-amended soils. The Bt genetic modification increased CO₂ production from stem-amended soils (P < 0.05) and decreased N mineralization in root-amended soils. The ¹³C and ¹⁵N results also showed more residue-C and -N retained in soils mixed with NBt stem residues. After 36 weeks leaf- and stem-amended soils with indulin lignin had a lower Ad/Al ratio and were less degraded than soils without exogenous lignin. In conclusion, plant lignin and nitrogen contents were good predictors of CO₂ production and N mineralization potential. Corn roots decomposed more slowly than aboveground components emphasizing the importance of recalcitrant root residues in sustaining the organic matter content of soil.     

Phenol oxidase, peroxidase and organic matter dynamics of soil

Extracellular enzymes mediate the degradation, transformation and mineralization of soil organic matter. The activity of cellulases, phosphatases and other hydrolases has received extensive study and in many cases stoichiometric relationships and responses to disturbances are well established. In contrast, phenol oxidase and peroxidase activities, which are often uncorrelated with hydrolase activities, have been measured in only a small subset of soil enzyme studies. These enzymes are expressed for a variety of purposes including ontogeny, defense and the acquisition of carbon and nitrogen. Through excretion or lysis, these enzymes enter the environment where their aggegrate activity mediates key ecosystem functions of lignin degradation, humification, carbon mineralization and dissolved organic carbon export. Phenol oxidases and peroxidases are less stable in the environment than extracellular hydrolases, especially when associated with organic particles. Activities are also affected, positively and negatively, by interaction with mineral surfaces. High spatiotemporal variation obscures their relationships with environmental variables and ecological process. Across ecosystems, phenol oxidase and peroxidase activities generally increase with soil pH, a finding not predicted from the pH optima of purified enzymes. Activities associated with plant litter and particulate organic matter often correlate with decomposition rates and potential activities generally increase with the lignin and secondary compound content of the material. At the ecosystem scale, nitrogen amendment alters the expression of phenol oxidase and peroxidase enzymes more broadly than culture studies imply and these responses correlate with positive and negative changes in litter decomposition rates and soil organic matter content. At the global scale, N amendment of basidiomycete-dominated soils of temperate and boreal forest ecoystems often leads to losses of oxidative enzyme activity, while activities in grassland soils dominated by glomeromycota and ascomycetes show little net response. Land use that leads to loss of soil organic matter tends to increase oxidative activities. Across ecosystems, soil organic matter content is not correlated with mean potential phenol oxidase and peroxidase activities. A multiple regression model that includes soil pH, mean annual temperature, mean annual precipitation and potential phenol oxidase activity accounts for 37% of the variation in soil organic matter (SOM) content across ecosystems (n = 63); a similar model for peroxidase activity describes 32% of SOM variance (n = 43). Analysis of residual variation suggest that suites of interacting factors create both positive and negative feedbacks on soil organic matter storage. Soils with high oxygen availability, pH and mineral activity tend to be substrate limited: high in situ oxidative activities limit soil organic matter accumulation. Soils with opposing characteristics are activity limited: low in situ oxidative activities promote soil organic matter storage.     

Microbial enzyme activities in leaf litter, humus and mineral soil layers of European forests

The rationale of the study was to investigate microbial activity in different soil horizons in European forests. Hence, activities of chitinase and cellulase, microbial biomass carbon (C_mic) and basal respiration were measured in litter, fragmentation, humus and mineral soil layers collected several times from various beech and spruce forests. Sites were selected to form a gradient in N availability. Analyses were also performed on beech litter from a litterbag transplant experiment. Furthermore, microbiological parameters were measured in horizons of beech and spruce chronosequence sites with different stand age in order to investigate the influence of forest rotation, and hence changes in soil organic matter (SOM) dynamics, on microbial activity. Finally in horizons of one beech forest, the seasonal variation of selected microbiological parameters was measured more intensively. β-Glucosaminidase and cellobiohydrolase activities were measured using fluorogenic 4-methylumbelliferyl substrates to estimate chitinase and cellulase activities, respectively. On a spatial scale, chitinase and cellulase activities, C_mic determined by substrate induced respiration, and basal respiration ranged from 144 to 1924 and 6-177 nmol 4-MU g⁻¹ org-C h⁻¹, 8-48 mg C g⁻¹ org-C and 11-149 μg CO₂-C g⁻¹ org-C h⁻¹, respectively; in general values were significantly lower in layers of humus and mineral soil than of litter. Chitinase activity, C_mic and basal respiration from humus and mineral soil layers, together, correlated positively, while none correlated with cellulase activity. Similarly in the litter layer, no correlations were found between the microbiological parameters. On a seasonal scale, a time lag between a burst in basal respiration rate and activities of both enzymes were observed. In general, activities of cellulase and chitinase, C_mic and basal respiration, did not change with stand age, except in the humus layer in the spruce chronosequence, where C_mic decreased with stand age. In the litter layer, cellulase activity was significantly and positively related to the C:N ratio, while only a tendency for chitinase activity was shown, indicating that enzyme activities decreased with increasing N availability. In accordance, the enzyme activities and C_mic decreased significantly with increasing chronic N deposition in the humus layer, while basal respiration only tended to decrease with increasing N deposition. In contrast, enzyme activities in beech litter from litterbags after 2 years of incubation were generally higher at sites with higher N deposition. The results show different layer-specific responses of enzyme activities to changes in N availability, indicating different impacts of N availability on decomposition of SOM and stage of litter decomposition.     

Interactions between litter quality, decomposition and soil fertility: a laboratory study

Leaf litters from beech (Fagus sylvatica L.) and oak (Quercus robur L.) trees were collected from mixed, deciduous woodlands growing on three soil types that varied in mineral nutrient concentrations and N mineralisation potential. Litter quality, including %N, %Mn, %P, acid detergent fibre, cellulose, Klason lignin, phenylpropanoid constituents of lignin, hexose and pentose sugar (mainly from hemicelluloses) varied within species according to soil type. However, oak and beech showed the opposite responses to soil nutrient status for most of these variables. The litters were incubated in the laboratory for 12 months (at 18 °C and constant moisture) on beds of forest floor material from two soils of contrasting high nutrient material (HNM) or low nutrient material (LNM) nutrient status to investigate litter quality and substrate interactions. At 4, 8 and 12 months there were significant differences in mass losses from oak and beech litters from all sites, and for each litter type exposed to the HNM and LMN soils. At 12 months mean mass losses were higher for HNM treatment (38.7% oak, 27.8% beech) than for the LNM treatment (30.6% oak, 25.5% beech). However, the beech and oak litters from the different sites consistently responded in opposite ways on the same soil treatment reflecting site-related effects on litter quality. Initial concentration of Klason lignin was the best predictor for mass losses from litter species and litter types. Intra-specific variation in rates of litter decomposition of beech and oak litters from different sites, and differences in their interactions with the two forest floor materials, illustrate the complexities of proximate controls on decomposition that are often masked in system-level studies.     

Location and chemical composition of stabilized organic carbon in topsoil and subsoil horizons of two acid forest soils

The ¹⁴C age of soil organic matter is known to increase with soil depth. Therefore, the aim of this study was to examine the stabilization of carbon compounds in the entire soil profile using particle size fractionation to distinguish SOM pools with different turnover rates. Samples were taken from a Dystric Cambisol and a Haplic Podzol under forest, which are representative soil types under humid climate conditions. The conceptual approach included the analyses of particle size fractions of all mineral soil horizons for elemental composition and chemical structure of the organic matter by ¹³C cross-polarization magic angle spinning nuclear magnetic resonance (CPMAS NMR) spectroscopy. The contribution of phenols and hydroxyalkanoic acids, which represent recalcitrant plant litter compounds, was analyzed after CuO oxidation.In the Dystric Cambisol, the highest carbon concentration as well as the highest percentage of total organic carbon are found in the <6.3 μm fractions of the B and C horizons. In the Haplic Podzol, carbon distribution among the particle size fractions of the Bh and Bvs horizons is influenced by the adsorption of dissolved organic matter. A relationship between the carbon enrichment in fractions <6.3 μm and the ¹⁴C activity of the bulk soil indicates that stabilization of SOM occurs in fine particle size fractions of both soils. ¹³C CPMAS NMR spectroscopy shows that a high concentration of alkyl carbon is present in the fine particle size fractions of the B horizons of the Dystric Cambisol. Decreasing contribution of O-alkyl and aromatic carbon with particle size as well as soil depth indicates that these compounds are not stabilized in the Dystric Cambisol. These results are in accordance with data obtained by wet chemical analyses showing that cutin/suberin-derived hydroxyalkanoic acids are preserved in the fine particle size fractions of the B horizons. The organic matter composition in particle size fractions of the top- and subsoil horizons of the Haplic Podzol shows that this soil is acting like a chromatographic system preserving insoluble alkyl carbon in the fine particle size fractions of the A horizon. Small molecules, most probably organic acids, dominate in the fine particle size fractions of the C horizons, where they are stabilized in clay-sized fractions most likely due to the interaction with the mineral phase. The characterization of lignin-derived phenols indicated, in accordance with the NMR measurements, that these compounds are not stabilized in the mineral soil horizons.