Fungal colonization as affected by litter depth and decomposition stage of needle litter

The present study was designated to evaluate the relative effects of litter depth and decomposition stage of needles on fungal colonization of needle litter in field experiments. The experiment was carried out in coniferous temperate forests in central Japan. Needle litter of Chamaecyparis obtusa and Pinus pentaphylla var. himekomatsu at two decomposition stages (recently dead and partly decomposed) were placed into the organic layer at two depths (on the surface of and beneath the litter layer). Fungal colonization of needles after 1 year was examined in terms of hyphal abundance and frequency of fungal species. Total and live hyphal length on needles were affected by the litter depth and (or) the decomposition stage of needles. Length of darkly pigmented hyphae on needles was 1.7-2.6 times greater beneath the litter layer than on the litter surface regardless of the decomposition stage of needles. Length of clamp-bearing hyphae in Pinus pentaphylla was 5.0-5.2 times greater in partly decomposed needles than in recently dead needles regardless of the litter depth. Frequencies of Pestalotiopsis spp. and Cladosporium cladosporioides were higher on recently dead needles than on partly decomposed needles and (or) were higher on the litter surface than beneath the litter layer. Frequencies of Trichoderma, Penicillium, and Umbelopsis species generally were higher on partly decomposed needles than on recently dead needles and were higher beneath the litter layer than on the surface.     

Leaf litter is the main driver for changes in bacterial community structures in the rhizosphere of ash and beech

The rhizosphere and the surrounding soil harbor an enormous microbial diversity and a specific community structure, generated by the interaction between plant roots and soil bacteria. The aim of this study was to address the influences of tree species, tree species diversity and leaf litter on soil bacterial diversity and community composition. Therefore, mesocosm experiments using beech, ash, lime, maple and hornbeam were established in 2006, and sampled in October 2008 and June 2009. Mesocosms were planted with one, three or five different tree species and treated with or without litter overlay.Cluster analysis of DGGE-derived patterns revealed a clustering of 2008 sampled litter treatments in two separated clusters. The corresponding treatments sampled in 2009 showed separation in one cluster. PCA analysis based on the relative abundance of active proteobacterial classes and other phyla in beech and ash single-tree species mesocosm indicated an effect of sampling time and leaf litter on active bacterial community composition. The abundance of next-generation sequencing-derived sequences assigned to the Betaproteobacteria was higher in the litter treatments, indicating a higher activity, under these conditions. The Deltaproteobacteria, Nitrospira and Gemmatimonadetes showed an opposite trend and were more active in the mesocosms without litter. The abundance of alphaproteobacterial sequences was higher in mesocosms sampled in 2009 (P = 0.014), whereas the Acidobacteria were more active in 2008 (P = 0.014). At the family level, we found significant differences of the litter vs. non-litter treated group. Additionally, an impact of beech and ash as tree species on soil bacterial diversity was confirmed by the Shannon and Simpson indices. Our results suggest that leaf litter decomposition in pH-stable soils affect the soil bacterial composition, while tree species influence the soil bacterial diversity.     

Responses of soil respiration,soil nutrients,and litter decomposition to inputs from canopy herbivores

We tested whether inputs from canopy herbivores would affect soil processes such as respiration, nutrient cycling, and decomposition along an elevation gradient. The five treatments we used were frass additions, throughfall additions, removal of all litter that fell during the study, removal of greenfall that fell during the study, and controls. Soil respiration was significantly reduced on low and mid elevation sites in litter exclusion, greenfall exclusion and throughfall addition treatments (from 0.846 g CO₂/m²/h for controls to 0.618, 0.667, and 0.708 g CO₂/m²/h, respectively, for the three treatments). Throughfall additions containing PO₄ and NH₄ contributed to significant increases in PO₄ (as much as 0.737 mg/l in 100 ml KCl extract greater then controls), but decreases in NO₃, (0.306  mg/l in 100 ml KCl extract less than con trols), in soil solution samples compared to controls. We observed no significant treatment effects on litter decomposition. Precipitation and temperature influenced soil respiration, but both factors showed a significant interaction with elevation. Phosphate concentrations in soil solutions differed significantly with elevation (low elevation mean 0.097 mg/l, mid elevation mean 0.192 mg/l). Elevation had no significant effect on decomposition.     

Animal manure and anhydrous ammonia amendment alter microbial carbon use efficiency,microbial biomass,and activities of dehydrogenase and amidohydrolases in semiarid agroecosystems

The potential negative impact of agricultural practices on soil and water quality is of environmental concern. The associated nutrient transformations and movements that lead to environmental concerns are inseparable from microbial and biochemical activities. Therefore, biochemical and microbiological parameters directing nitrogen (N) transformations in soils amended with different animal manures or inorganic N fertilizers were investigated. Soils under continuous corn cultivation were treated with N annually for 5 years at 56, 168, and 504 kg N ha⁻¹ in the form of swine effluent, beef manure, or anhydrous ammonia. Animal manure treatments increased dehydrogenase activity, microbial biomass carbon (C_mic) and N (N_mic) contents, and activities of amidohydrolases, including l-asparaginase, urease, l-glutaminase, amidase, and β-glucosaminidase. Soils receiving anhydrous ammonia demonstrated increased nitrate contents, but reduced microbiological and biochemical activities. All treatments decreased C_mic:organic C (C_org) ratios compared with the control, indicating reduced microbial C use efficiency and disturbance of C equilibrium in these soil environments. Activities of all enzymes tested were significantly correlated with soil C_org contents (P < 0.001, n = 108), but little correlation (r = 0.03, n = 36) was detected between C_mic and C_org. Activities of amidase and β-glucosaminidase were dominated by accumulated enzymes that were free of microbial cells, while activities of asparaginase and glutaminase were originated predominately from intracellular enzymes. Results indicated that soil microbial and biochemical activities are sensitive indicators of processes involved in N flow and C use efficiency in semiarid agroecosystems.     

Dynamics of litter carbon turnover and microbial abundance in a rye detritusphere

Factors determining C turnover and microbial succession at the small scale are crucial for understanding C cycling in soils. We performed a microcosm experiment to study how soil moisture affects temporal patterns of C turnover in the detritusphere. Four treatments were applied to small soil cores with two different water contents (matric potential of ?0.0063 and ?0.0316 MPa) and with or without addition of ¹³C labelled rye residues (δ¹³C=299‰), which were placed on top. Microcosms were sampled after 3, 7, 14, 28, 56 and 84 days and soil cores were separated into layers with increasing distance to the litter. Gradients in soil organic carbon, dissolved organic carbon, extracellular enzyme activity and microbial biomass were detected over a distance of 3 mm from the litter layer. At the end of the incubation, 35.6% of litter C remained on the surface of soils at ?0.0063 MPa, whereas 41.7% remained on soils at ?0.0316 MPa. Most of the lost litter C was mineralised to CO₂, with 47.9% and 43.4% at ?0.0063 and ?0.0316 MPa, respectively. In both treatments about 6% were detected as newly formed soil organic carbon. During the initial phase of litter decomposition, bacteria dominated the mineralisation of easily available litter substrates. After 14 days fungi depolymerised more complex litter compounds, thereby producing new soluble substrates, which diffused into the soil. This pattern of differential substrate usage was paralleled by a lag phase of 3 days and a subsequent increase in enzyme activities. Increased soil water content accelerated the transport of soluble substrates, which influenced the temporal patterns of microbial growth and activity. Our results underline the importance of considering the interaction of soil microorganisms and physical processes at the small scale for the understanding of C cycling in soils.     

Effect of litter quality and soil fungi on macroaggregate dynamics and associated partitioning of litter carbon and nitrogen

We investigated the effect of plant residue decomposability and fungal biomass on the dynamics of macroaggregate (250–2000 μm) formation in a three months' incubation experiment and determined the distribution of residue-derived C and N in the microbial biomass and in aggregate size fractions (250–2000 μm, 53–250 μm and <53 μm) using ¹³C and ¹⁵N data. A silty loam soil (sieved <250 μm) was incubated with and without addition of ¹⁵N labelled maize leaves (C/N = 27.4) and roots (C/N = 86.4). Each treatment was carried out with and without fungicide application. The addition of maize residues enhanced soil respiration and microbial biomass C and N and resulted in increased macroaggregate formation with a higher and more rapid maximum macroaggregation in the soil amended with maize leaves than in that with addition of roots. Fungicide application led to a significant decline of microbial biomass C and mineralization of the added residues compared to untreated soils, which demonstrates a successful suppression of part of the active microbial biomass by the fungicide. However, this was not confirmed by a generally lower ergosterol concentration. Consequently, ergosterol was no reliable fungal biomarker in periods of rapid decline of the fungal biomass. A single addition of fungicide was insufficient for continued inhibition of the fungal biomass. Yet, a significant delay (28–42 days) in macroaggregation in fungicide treated compared to untreated samples highlighted the importance of the fungal biomass in macroaggregate formation. Macroaggregates were enriched in maize-derived ¹³C and ¹⁵N compared to microaggregates or the fraction < 53 μm. They turned over rapidly with decreasing substrate availability, which entailed a transfer of maize-derived C and N stored within macroaggregates during the first weeks of incubation to microaggregates with proceeding incubation time. Our results indicate that this transfer happened within macroaggregates, because no considerable amount of free particulate organic matter (POM) was released upon macroaggregate breakdown. We conclude that substrate decomposability and fungal activity are key factors determining extent and dynamics of macroaggregation during decomposition processes. Macroaggregate formation implied rapid incorporation and thereby short-term protection of maize-derived C and N. Moreover, macroaggregates allowed a transfer of maize-derived organic matter into microaggregates within macroaggregates, which prevented the release of significant amounts of free POM upon macroaggregate breakdown. Consequently, macroaggregates constitute to the transfer of recently added C into more stable soil organic matter fractions.     

Carbon flow into microbial and fungal biomass as a basis for the belowground food web of agroecosystems

The origin and quantity of plant inputs to soil are primary factors controlling the size and structure of the soil microbial community. The present study aimed to elucidate and quantify the carbon (C) flow from both root and shoot litter residues into soil organic, extractable, microbial and fungal C pools. Using the shift in C stable isotope values associated with replacing C3 by C4 plants we followed root- vs. shoot litter-derived C resources into different soil C pools. We established the following treatments: Corn Maize (CM), Fodder Maize (FM), Wheat + maize Litter (WL) and Wheat (W) as reference. The Corn Maize treatment provided root- as well as shoot litter-derived C (without corn cobs) whereas Fodder Maize (FM) provided only root-derived C (aboveground shoot material was removed). Maize shoot litter was applied on the Wheat + maize Litter (WL) plots to trace the incorporation of C4 litter C into soil microorganisms. Soil samples were taken three times per year (summer, autumn, winter) over two growing seasons. Maize-derived C signal was detectable after three to six months in the following pools: soil organic C (C_org), extractable organic C (EOC), microbial biomass (C_mic) and fungal biomass (ergosterol). In spite of the lower amounts of root- than of shoot litter-derived C inputs, similar amounts were incorporated into each of the C pools in the FM and WL treatments, indicating greater importance of the root- than shoot litter-derived resources for the soil microorganisms as a basis for the belowground food web. In the CM plots twice as much maize-derived C was incorporated into the pools. After two years, maize-derived C in the CM treatment contributed 14.1, 24.7, 46.6 and 76.2% to C_org, EOC, C_mic and ergosterol pools, respectively. Fungi incorporated maize-derived C to a greater extent than did total soil microbial biomass.     

Grasses, litter, and their interaction affect microbial biomass and soil enzyme activity

Plant effects on ecosystem processes are mediated through plant-microbial interactions belowground and soil enzyme assays are commonly used to directly relate microbial activity to ecosystem processes. Live plants influence microbial biomass and activity via differences in rhizosphere processes and detrital inputs. I utilized six grass species of varying litter chemistry in a factorial greenhouse experiment to evaluate the relative effect of live plants and detrital inputs on substrate-induced respiration (SIR, a measure of active microbial biomass), basal respiration, dissolved organic carbon (DOC), and the activities of β-glucosidase, β-glucosaminidase, and acid phosphatase. To minimize confounding variables, I used organic-free potting media, held soil moisture constant, and fertilized weekly. SIR and enzyme activities were 2-15 times greater in litter-addition than plant-addition treatments. Combining live plants with litter did not stimulate microbial biomass or activity above that in litter-only treatments, and β-glucosidase activity was significantly lower. Species-specific differences in litter N (%) and plant biomass were related to differences in β-glucosaminidase and acid phosphatase activity, respectively, but had no apparent effect on β-glucosidase, SIR, or basal respiration. DOC was negatively related to litter C:N, and positively related to plant biomass. Species identity and living plants were not as important as litter additions in stimulating microbial activity, suggesting that plant effects on soil enzymatic activity were driven primarily by detrital inputs, although the strength of litter effects may be moderated by the effect of growing plants.     

Soil fungal population levels in cotton fields fertilized with poultry litter and their relationships to soil nutrient concentrations and plant growth parameters

Application to land of large quantities of waste materials from concentrated animal production, without causing environmental pollution, presents a major challenge to agriculture in the 21st century. Effects of land-applied animal wastes on chemical contents of soil are well documented, but less is understood of their effects on microbial populations in soil. This study was undertaken to evaluate effects of commercial application of poultry litter (PL), as a fertilizer, on soil fungal population levels and components on cotton farms in Mississippi, and to determine relationships of fungal population levels to soil nutrient contents and cotton growth and yield. On each of two farms, soil fungal population levels were estimated by dilution plating from samples of soil collected at two sampling times during 2 years from replicated plots of four fertilization treatments: 0 fertilizer, conventional mineral fertilizer (CF), low PL, and high PL. Soil fungal population levels differed significantly (P = 0.05) according to years or seasons and fertilization treatments on both farms. Population levels often were higher in soils amended with low or high PL, or with CF, than in unfertilized controls. On one farm where PL was tilled into soil, fungal population levels increased significantly during the course of the experiment in PL and CF treatments, but not in unfertilized controls, as determined by linear regression. No such increases were observed on the second farm where PL was applied no-till. Population levels of Fusarium semitectum and Penicillium purpurogenum were significantly higher in PL-treated soils than in unfertilized controls in 1 or 2 of 4 sampling events at both farms, while levels of four other species or groups of fungi usually did not differ. Fungal population levels were significantly correlated (P = 0.05) with N concentrations of soils in 1–4 sampling events on each farm and less frequently correlated with concentrations of nine other elements. In 4 of 24 instances, soil fungal population levels were significantly correlated with leaf area index, chlorophyll content, or yield of cotton, and correlation coefficients with these plant parameters were always positive. Major conclusions derived from this study are (1) no deleterious effects on population levels of total or select soil fungi were observed with use of PL as a fertilizer for commercial cotton production; and (2) soil fungal population levels may increase over time in association with greater fertility and plant growth that is induced by both mineral fertilizer and PL applications.     

Low levels of nitrogen addition stimulate decomposition by boreal forest fungi

Climate warming and associated increases in nutrient mineralization may increase the availability of soil nitrogen (N) in high latitude ecosystems, such as boreal forests. These changes in N availability could feed back to affect the decomposition of litter and organic matter by soil microbes. Since fungi are important decomposers in boreal forest ecosystems, we conducted a 69-day incubation study to examine N constraints on fungal decomposition of organic substrates common in boreal ecosystems, including cellulose, lignin, spruce wood, spruce needle litter, and moss litter. We added 0, 20, or 200 μg N to vials containing 200 mg substrate in factorial combination with five fungal species isolated from boreal soil, including an Ascomycete, a Zygomycete, and three Basidiomycetes. We hypothesized that N addition would increase CO₂ mineralization from the substrates, particularly those with low N concentrations. In addition we predicted that Basidiomycetes would be more effective decomposers than the other fungi, but would respond weakly or negatively to N additions. In support of the first hypothesis, cumulative CO₂ mineralization increased from 635 ± 117 to 806 + 108 μg C across all fungal species and substrates in response to 20 μg added N; however, there was no significant increase at the highest level of N addition. The positive effect of N addition was only significant on cellulose and wood substrates which contained very little N. We also observed clear differences in the substrate preferences of the fungal species. The Zygomycete mineralized little CO₂ from any of the substrates, while the Basidiomycetes mineralized all of the substrates except spruce needles. However, the Ascomycete (Penicillium) was surprisingly efficient at mineralizing spruce wood and was the only species that substantially mineralized spruce litter. The activities of β-glucosidase and N-acetyl-glucosaminidase were strongly correlated with cumulative respiration (r = 0.78 and 0.74, respectively), and Penicillium was particularly effective at producing these enzymes. On moss litter, the different fungal species produced enzymes that targeted different chemical components. Overall, our results suggest that fungal species specialize on different organic substrates, and only respond to N addition on low N substrates, such as wood. Furthermore, the response to N addition is non-linear, with the greatest substrate mineralization at intermediate N levels.     

Changes in litter properties during decomposition: A study by differential thermogravimetry and scanning calorimetry

To verify the paradigm that organic matter (OM) quality (q) decreases with decomposition it is necessary to define q in strictly chemical, operational terms. We suggest defining q as the result of a balance between the energy stored in OM and the external supply of energy needed to release it. We apply this concept to the study of litter decomposition in four European pine forests: boreal, cool Atlantic, Mediterranean and warm Atlantic. Intact litter cores were taken and transported to the laboratory, where needles were sorted into six classes that summarize the main facts of the decomposition: melanisation, fragmentation and perforation by mesofauna. Each class was analyzed by both differential thermogravimetry and differential scanning calorimetry to obtain its spectra of weight loss and energy release.In the non-decomposed needles, two peaks of weight loss and energy release appear: a labile peak at about 350 °C, and a recalcitrant peak at about 450 °C. During decomposition, both peaks (but especially the recalcitrant one) move to lower temperatures, and their shapes change from well defined to flattened. In Mediterranean litters, a third peak appears at about 500 °C, due probably to refractory products of neoformation. There is a continuous increase in the energy stored in the remaining litter (in Joules per unit OM): this increase is concentrated in both the most thermolabile fractions (lost at temperatures <350 °C) and the most thermostable ones (>450 °C). With decomposition OM becomes more recalcitrant (i.e., it is lost at higher temperatures), but its stored energy becomes more available (i.e., it is released at lower temperatures). Overall, the energetic benefit/cost ratio increases. Thus, our results to date do not agree with the current paradigm that q decreases with decomposition; rather, they suggest that, at least in the first phases we studied, q is maintained or even increases.     

Photoacceleration of plant litter decomposition in an arid environment

In arid ecosystems, abiotic processes facilitate the physical and chemical degradation of plant litter to the extent that decomposition models that use climatic and litter composition variables as surrogates for microbial activity are not predictive. The purpose of this study was to estimate the potential contribution of photodegradation to the decomposition of plant litters that varies in architecture and chemical composition. Litter of Pinus edulis, Juniperus monosperma and Populus deltoides were exposed to ambient and attenuated sunlight, with and without supplemental water additions, at a riparian forest site along the Middle Rio Grande (New Mexico, USA). Mass loss, elemental composition, and microbial extracellular enzyme activities (EEA) were measured over 639 days. The composition of the fungal communities associated with the decomposing litters was compared by analyses of fungal ITS nrDNA sequences. Litter exposed to ambient sunlight had greater mass loss rates than shaded litter, independent of the water treatment: Populus increased by 100%, Pinus by 86% and Juniperus by 46%. The increases were proportional to exposed litter surface area per g dry mass. EEA potentials, particularly oxidative activities, were low in comparison to those measured in mesic ecosystems. For Populus litter, the principal driver of photoacceleration appeared to be photodegradation of cellulose; for Pinus, it was photodegradation of polyphenols; for Juniperus accelerated mass loss was associated with photodegradation of both polysaccharides and polyphenols. Fungal community composition varied by litter type, but the dominant colonizers were yeasts and dark-septate hyphal taxa; a finding consistent with the low enzymatic oxidation potential. This study shows that photochemical oxidation can supplement enzymatic oxidation and increase decomposition rates. As a result, organic matter decomposition in arid ecosystems is not restricted to periods of high moisture availability as is plant production. This decoupling may partly account for the low soil organic matter content of these ecosystems.     

Soil compaction and poultry litter effects on factors affecting nitrogen availability in a claypan soil

Soil compaction may affect N mineralization and the subsequent fate of N in agroecosystems. Laboratory incubation and field experiments were conducted to determine the effects of surface soil compaction on soil N mineralization in a claypan soil amended with poultry litter (i.e., Turkey excrement mixed with pine shavings as bedding). In a laboratory study, soil from the surface horizon of a Mexico silt loam soil was compacted to four bulk density levels (1.2, 1.4, 1.6 and 1.8 Mg m⁻³) with and without poultry litter and incubated at 25 °C for 42 days. A field trial planted to corn (Zea mays L.) was also conducted in 2002 on a Mexico silt loam claypan soil in North Central Missouri. Soil was amended with litter (0 and 19 Mg ha⁻¹) and left uncompacted or uniformly compacted. Soil compaction decreased soil inorganic N by a maximum of 1.8 times in the laboratory study; this effect was also observed at all depths of the field trial. Compacted soil with a litter amendment accumulated NH₄⁺-N up to 7.2 times higher than the noncompacted, litter-amended soil until Day 28 of the laboratory incubation and in the beginning of the growing season of the field study. Ammonium accumulation may have been due to decreased soil aeration under compacted conditions. Application of litter increased soil N mineralization throughout the growing season. In the laboratory study, soil inorganic N in unamended soil was negatively correlated with soil bulk density and the proportion of soil micropores, but was positively related with soil total porosity and the proportion of soil macropores. These results indicate that soil compaction, litter application and climate are interrelated in their influences on soil N mineralization in agroecosystems.     

Effects of NH4+ and NO3− on litter and soil organic carbon decomposition in a Chinese fir plantation forest in South China

Soil organic carbon (SOC) dynamics and nutrient availability determine the soil quality and fertility in a Chinese fir plantation forest in subtropical China. Uniformly ¹³C-labeled Chinese fir (Cunninghamia lanceolata) and alder (Alnus cremastogyne) leaf litter with or without 100 mg NH₄⁺ or NO₃⁻ were added to the soil. The purpose was to investigate the influence of N availability on the decomposition of the litter and native SOC. The production of CO₂, the natural abundance of ¹³C–CO₂, and the inorganic N dynamics were monitored. The results showed that Chinese fir (with a high C:N ratio) and alder (with a low C:N ratio) leaf litter caused significant positive priming effects (PEs) of 24% and 42%, respectively, at the end of the experiment (235 d). The PE dynamics showed that positive PE can last for at least 87 d. However, the possible occurrence of a significant negative PE with a sufficient incubation period is difficult to confirm. The application of both NH₄⁺ and NO₃⁻ was found to have a stimulating effect on the decomposition of Chinese fir and alder leaf litter in the early stage (0–15 d) of incubation, but an adverse effect in the late stage. Compared with NO₃⁻, NH₄⁺ caused a greater decrease in the PE induced by both Chinese fir and alder leaf litter. The effects of NH₄⁺ and NO₃⁻ on the PE dynamics had different patterns for different incubation stages. This result may indicate that the stability or recalcitrance of SOC, especially in such plantation forest soils, strongly depends on available leaf litter and application of N to the soil.     

Effects of fungal inocula on the decomposition of lignin and structural polysaccharides in Pinus sylvestris litter

 Litter bags containing sterile Scots pine (Pinus sylvestris) needles (19.8% lignin, 26.5% cellulose and 0.34% N) were inoculated with two species of fungi in the laboratory and then placed in the litter layer of a pine plantation. Marasmius androsaceus, which can degrade lignocellulose, was initially displaced by other fungal colonisers and was not detected in the litter after 2–3 months; but was re-isolated from the needles after 12 months. Trichoderma viride, which is a cellulolytic species and also antagonistic to other fungi, dominated the litter throughout the experiment. The control litter was naturally colonised by litter fungi. After 12 months, mass losses were similar at 52% for M. androsaceus and 48% for T. viride, compared with 36% for the control litter colonised by a more complex fungal community. Lignin concentrations increased with time in control litter and with T. viride because mass losses of carbohydrates were greater than those of lignin. Litter inoculated with M. androsaceus showed significant lignin decomposition throughout the experiment but cellulose concentrations showed a proportional increase in the first 6 months, suggesting that the fungus was preferentially exploiting hemicellulose and non-structural carbohydrates. Analysis of TFA-extractable sugars (mainly from hemicellulose) and CuO-derived phenylpropanoid moieties from lignin confirmed the differential patterns of resource decomposition which were not evident from total mass losses. During the initial stages of decomposition, T. viride was as effective in utilising structural polysaccharides as the complex fungal community in the control litter. Furthermore, M. androsaceus not only exhibited unexpectedly low cellulolytic activity but also facilitated lignin depolymerisation after the fungus was no longer detectable in the litter. The pre-inoculation of litter with these two fungal species therefore affected the overall dynamics of decomposition at a biochemical level. This study illustrates the importance of understanding the effects and interactions of specific fungi, rather than assumptions about the functional competence of diverse communities, on the processes of litter decomposition. Received: 5 July 2000     

Direct and indirect effects of nitrogen deposition on litter decomposition

Elevated nitrogen (N) deposition can affect litter decomposition directly, by raising soil N availability and the quantity and quality of litter inputs, and indirectly by altering plant community composition. We investigated the importance of these controls on litter decomposition using litter bags placed in annual herb based microcosm ecosystems that had been subject to two rates of N deposition (which raised soil inorganic N availability and stimulated litter inputs) and two planting regimes, namely the plant species compositions of low and high N deposition environments. In each microcosm, we harvested litter bags of 10 annual plant species, over an 8-week period, to determine mass loss from decomposition. Our data showed that species differed greatly in their decomposability, but that these differences were unlikely to affect decomposition at the ecosystem level because there was no correlation between a species’ decomposability and its response to N deposition (measured as population seed production under high N, relative to low N, deposition). Litter mass loss was ～2% greater in high N deposition microcosms. Using a comprehensive set of measurements of the microcosm soil environments, we found that the most statistically likely explanation for this effect was increased soil enzyme activity (cellobiosidase, β-glucosidase and β-xylosidase), which appears to have occurred in response to a combination of raised soil inorganic N availability and stimulated litter inputs. Our data indicate that direct effects of N deposition on litter input and soil N availability significantly affected decomposition but indirect effects did not. We argue that indirect effects of changes to plant species composition could be stronger in natural ecosystems, which often contain a greater diversity of plant functional types than those considered here.     

Combined effects of zinc and earthworm density on soil ecosystem functioning

In traditional environmental risk assessment for soils, interactions between biota, contaminants and soil functioning are seldom taken into account. Also, single species toxicity tests are conducted with a fixed number of test animals. The objective of this study was to investigate effects of zinc (0–620 mg Zn kg^?1 dry soil) on soil ecosystem processes at different densities of the earthworm Lumbricus rubellus. Experiments were conducted using 1-liter microcosms equipped with respirometers. The presence of L. rubellus stimulated relevant soil processes and parameters: litter fragmentation, leaf litter mass loss from the soil surface, soil organic matter (SOM) content and soil respiration. Zinc was not lethal to L. rubellus, but negatively impacted soil respiration at the highest concentrations. Litter mass loss from the soil surface was also decreased by zinc and there was a significant interaction with worm density. The results of the study demonstrate that the impact of zinc on soil processes depends on the presence and densities of key soil organisms such as earthworms that influence decomposition and SOM content. The outcome of this research can be used to make existing models for site-specific risk assessment more ecologically relevant, linking effects of contaminants on soil fauna populations with effects on ecosystem functioning.     

Effects of UV-B filtration on the chemistry and decomposition of Fraxinus excelsior leaves

Several studies have demonstrated a range of effects of outdoor UV-B supplementation during the growing season on leaf chemistry including carbohydrate extractability and on the subsequent decomposition of leaf litter. However, this study investigates the effects of several levels of UV radiation on leaf carbohydrate chemistry and subsequent decomposition using filtration of ambient sunlight. Fraxinus excelsior seedlings were grown outdoors in the UK under ambient solar irradiation and under filtration treatments which excluded either UV-B or both UV-A and UV-B. After one year of decomposition in the litter layer of a mixed semi-natural woodland, the loss of dry mass was 10% greater, relative to starting mass, in the leaves which had received no UV at all or no UV-B throughout the growing season (P < 0.05). Analysis of the cell wall material before decomposition revealed no significant trends in total carbohydrate and lignin content with UV exclusions, no change in foliar nitrogen and C-to-N ratio and a 2% increase in foliar carbon (P < 0.05) only with the combined exclusion of UV-A and UV-B. A sequential extraction of carbohydrate with a series of extractants (phosphate buffer, ammonium oxalate, urea, sodium hydroxide and formic acid) showed no trends with UV exclusions but digestion with the fungal enzyme mixture Driselase revealed that exclusion of UV-B only caused rhamnose and mannose residues of the cell-wall polysaccharides to resist Driselase digestion whist exclusion of all UV had the opposite effect. Whereas some studies have reported that elevated UV-B radiation from lamp supplementation can increase rates of subsequent leaf decomposition, the higher UV-B levels in the ambient controls of this filtration study resulted in 29% lower decomposition rates than the filtered-UV treatments.     

Deciduous woodland exposed to elevated atmospheric CO2 has species-specific impacts on anecic earthworms

Elevated atmospheric CO₂ induced reductions in litter quality can adversely affect earthworms. However, this understanding is based on laboratory rather than field research and relates to single earthworm and tree species. Here earthworm populations were investigated under Alnus glutinosa, Betula pendula, and Fagus sylvatica in a Free Air Carbon dioxide Enrichment field experiment. Litters from this experiment were also fed to Lumbricus terrestris L. at two rates with live weight change and cast properties assessed. Elevated CO₂ (580 ppmv) reduced litter N (−12%) with a corresponding increase in C:N ratio, especially for A. glutinosa. In the field, elevated CO₂ caused a shift in overall population composition, mainly characterised by reduced anecic biomass (–25%); endogeic and epigeic species were less affected. CO₂ effects on total biomass were most pronounced for A. glutinosa (e.g. field total biomass −47% vs. −11% overall). Growth of L. terrestris was lower when fed elevated CO₂ litter (−18%), although increased inputs of A. glutinosa litter mitigated this effect. In mesocosms, fresh cast respiration was lower (−14%) for elevated CO₂ litter, an effect more pronounced for A. glutinosa (−24%). When normalised for C content, elevated CO₂ effects on cast respiration were again negative and most marked for A. glutinosa litter. Litter N concentration, and possibly ease of litter mineralisation were factors affecting litter resource quality Litter N and P concentrations varied with A. glutinosa > B. pendula > F. sylvatica; F. sylvatica had the highest cellulose content. Field earthworm biomass was higher under A. glutinosa compared with B. pendula and F. sylvatica (+17 and +70%, respectively); live weight increased with A. glutinosa litter in the feeding trial almost three times more than for B. pendula, whereas it decreased for F. sylvatica. Cast respiration was highest for A. glutinosa, intermediate for B. pendula (ca. −36%) and lowest for F. sylvatica (ca. −78%). Earthworm responses to elevated CO₂ were complex, being characteristic of individual tree and earthworm species; responses were more adverse for trees with higher quality litter and for anecic earthworms.     

Application of 13C-labeled litter and root materials for in situ decomposition studies using phospholipid fatty acids

Microorganisms play a central role in litter decomposition and partitioning C between CO₂ evolution and sequestration of C into semi-permanent pools in soils. At the ecosystem level, forest stand age influences rates of litter accumulation and quality, and micro-climatology which could affect the microbial community structure and C sequestration processes. Although numerous laboratory experiments have studied the decomposition of model ¹³C-labeled compounds, few studies have verified these findings under field conditions. The objective of this study was to track decomposition of ¹³C-labeled Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) materials into the soil microbial community using ¹³C-phospholipids fatty acid (PLFA) analysis in three different aged forest stands. A field experiment was conducted that had three forest stand age treatments: old-growth (>500 yrs); 8-year-old clear-cut (CC8); and 25-year-old clear-cut (CC25) (landscape reps of n = 2). Each stand age had in situ microcosms that were amended with either ¹³C-labeled surface litter or root material. Microcosms were destructively sampled seven times over a 22-month period and the soil was analyzed for the relative amounts of ¹³C incorporated (¹³C%_INCORP) into PLFAs and the proportional distribution of ¹³C incorporated into PLFAs. The ¹³C%_INCORP was affected by stand age and ¹³C source with greater ¹³C%_INCORP in samples from CC8 than OG or CC25. Also, the level of ¹³C%_INCORP was greater for labeled litter than root material in five out of the seven sample dates. In general, 18:1ω9 and 18:2ω6,9 (common fungal biomarkers) had the greatest amount of ¹³C incorporation throughout the study period in both clear-cut and old-growth sites, especially in plots with ¹³C-labeled litter. Our data showed a low fungal ¹³C-PLFA: bacterial ¹³C-PLFA ratio (0.45) 1 month after incubation was initiated compared to 5, 7 and 9 months after incubation (two of these dates were >1.0). This suggests that initially bacteria played a greater role in the decomposition of the added needles with fungi playing a more important role in subsequent sample dates. Our results illustrate that the use of ¹³C-labeled materials in field studies coupled with¹³C-PLFA profiling is a powerful tool for determining microbial dynamics during decomposition – enabling statistically significant detection of land management treatment effects on C acquisition by microbial functional groups.