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.     

Nutrient limitation of litter decomposition with long-term secondary succession: evidence from controlled laboratory experiments

Purpose

Understanding ecosystem processes such as litter decomposition in response to dramatic land-use change is critical for modeling and predicting carbon (C) cycles. However, the patterns of litter decomposition along with long-term secondary succession (over 100 years) are not well reported, especially concerning nutrient limitations on litter decomposition.

Materials and methods

To clarify the response of litter decomposition to changes in soil nutrient availability, we conducted four incubation experiments involving soil and litter and nutrient addition from different successional stages and investigated the changes in microbial respiration and litter mass loss.

Results and discussion

Our results revealed that microbial respiration increased with succession without any litter addition (1.19~1.73 mg C g^?1 soil), and litter addition significantly promoted microbial respiration (16.5~72.9%), especially in the early successional stage (grassland and shrubland). The decomposition rate of the same litter decreased with succession. In addition, nitrogen (N) and phosphorus (P) addition showed significant effects on litter decomposition and microbial respiration; P addition promoted litter decomposition (2.4~15.3%) and microbial respiration (10.1~34.5%) in all successional stages, while N addition promoted litter decomposition (4.0~10.3%) and microbial respiration (5.4~27.2%) in all except the last stage of succession, which showed a negative effect on litter decomposition (??7.5%) and microbial respiration (??6.1%), indicating possible N saturation of litter decomposition and microbial respiration.

Conclusions

This work highlights that soil nutrient availability and successional stages need to be taken into account to predict the changes to litter decomposition in response to global changes.

     

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].     

A litter-slurry technique elucidates the key role of enzyme production and microbial dynamics in temperature sensitivity of organic matter decomposition

The rate at which organic matter decomposes generally increases with temperature, unless it is physico-chemically protected from enzymatic depolymerization. The temperature sensitivity of decomposition should increase with decreasing reaction rates, corresponding to increasing activation energy of the decomposing compounds. One approach to testing this carbon-quality temperature hypothesis is to study the effect of temperature on leaf litter decomposition, because fresh surface litter is unprotected. However, other factors such as humidity co-vary with temperature, and biological processes such as enzyme production and microbial population growth may also be thermally sensitive. We developed a litter slurry approach to isolate the effect of temperature and litter quality on decomposition. We found that pine litter decomposed faster than oak litter, consistent with a lower C:N and lignin:N ratio. During the first 14 days of decomposition, there was no difference in decomposition rate for litter incubated at 25 °C compared to 35 °C. Lower potential enzyme activity at 35 °C suggested that enzyme production was suppressed at 35 °C compared to 25 °C, resulting in similar in situ enzyme activities at the two temperatures. After 14 days, enzyme pools were similar between the two incubation temperatures, which resulted in faster decomposition at the warmer temperature, consistent with enzyme kinetic theory. At Day 14, the decomposition rate of the high quality pine litter was more temperature sensitive than the decomposition rate of the lower quality oak litter, suggesting that the quality of soluble pool rather than bulk chemistry determined the temperature sensitivity during this stage. After 28 days of incubation, oak litter decomposition was more temperature sensitive than pine litter, consistent with the carbon temperature-quality hypothesis. The litter slurry approach revealed that biological responses to temperature can affect the apparent temperature sensitivity of decomposition, and highlight a need for further research into microbial responses to temperature.     

Microbial enzyme activities,fungal biomass and quality of the litter and upper soil layer in a beech forest of south Italy

Forest soils contain a large amount of organic matter (OM) and therefore represent a considerable carbon reserve. The amount of OM sequestered in the soil is dependent on annual input of litter and its quality. The aim of this study was to investigate the quantity and quality of OM, the microbial capacity to degrade it and its recalcitrance to further degradation, by considering some extracellular enzyme activities in a beech (Fagus sylvatica L.) forest in south Italy (Mediterranean area). Our attention was focused on the decomposition continuum of the litter horizon and upper soil layer. Because fungi are the major decomposers of plant material, fungal biomass was also measured and its relationship with enzyme activities was tested. The results showed that: (i) the litter horizon and the upper soil layer differed in chemical characteristics and biological activities; (ii) within the litter horizon, the three layers detected for their different degree of degradation (L, recently fallen, not decomposed and not compressed material; F, partially decomposed and fragmented but macroscopically recognizable material; H, compressed and strongly fragmented) differed more in chemical characteristics than in biological activities; (iii) the enzyme activities and fungal biomass changed during the study period but a clear relationship with succession of seasons was evident only for cellulase, laccase, peroxidase and fungal biomass; and (iv) the upper soil layer included 42% OM and less than 50% of that was susceptible to further decomposition. This percentage was 30% in the OM of L.     

The oribatid mite Scheloribates moestus (Acari: Oribatida) alters litter chemistry and nutrient cycling during decomposition

It is widely accepted that microarthropods influence decomposition dynamics but we know relatively little about their effects on litter chemistry, extracellular enzyme activities, and other finer-scale decomposition processes. Further, few studies have investigated the role of individual microarthropod species in litter decomposition. The oribatid mite Scheloribates moestus Banks (Acari: Oribatida) is abundant in many U.S. ecosystems. We examined the potential effects of S. moestus on litter decomposition dynamics and chemical transformations, and whether these effects are influenced by variation in initial litter quality. We collected corn and oak litter from habitats with large populations of S. moestus and in microcosms with and without mites measured respiration rates, nitrogen availability, enzyme activities, and molecular-scale changes in litter chemistry. Mites stimulated extracellular enzyme activities, enhanced microbial respiration rates by 19% in corn litter and 17% in oak litter over 62 days, and increased water-extractable organic C and N. Mites decreased the relative abundance of polysaccharides in decomposing corn litter but had no effect on oak litter chemistry, suggesting that the effects of S. moestus on litter chemistry are constrained by initial litter quality. We also compared the chemistry of mite feces to unprocessed corn litter and found that feces had a higher relative abundance of polysaccharides and phenols and a lower relative abundance of lignin. Our study establishes that S. moestus substantially changes litter chemistry during decomposition, but specific effects vary with initial litter quality. These chemical transformations, coupled with other observed changes in decomposition rates and nutrient cycling, indicate that S. moestus could play a key role in soil C cycling dynamics.     

Decomposition and mineralization of energy crop residues governed by earthworms

Energy crops are increasingly cultivated in agricultural management systems world-wide. A substitution of food crops (e.g. cereals) by energy crops may generally alter the biological activity and litter decomposition in soil due to their varying structural and chemical composition and subsequently modify soil functioning. A soil microcosm experiment was performed to assess the decomposition and microbial mineralization of different energy crop residues in soil compared to a food crop, with or without earthworms. Residues of the energy crops winter rape (Brassica napus), maize (Zea mays), miscanthus (Miscanthus giganteus) and the food crop oat (Avena sativa) were each provided as food source for a mixed earthworm population, each consisting of one individual of Lumbricus terrestris, Aporrectodea caliginosa, and Octolasion tyrtaeum. After 6 weeks, the rate of litter loss from the soil surface, earthworm biomass, microbial biomass-C and -N, microbial activity, and enzyme activities were determined. The results emphasized, that litter loss and microbial parameters were predominantly promoted by earthworms and were additionally influenced by the varying structural and chemical composition of the different litter. Litter decay by earthworms was highest in N-rich maize litter treatment (C-N ratio 34.8) and lowest in the case of miscanthus litter (C-N ratio 134.4). As a consequence, the microbial biomass and basal respiration in soils with maize litter were higher, relative to other litter types. MBC-MBN ratio in soil increased when earthworms were present, indicating N competition between earthworms and microorganisms. Furthermore, enzyme activities responded in different ways on the varying types of litter and earthworm activity. Enzymes involved in the N-cycle decreased and those involved in the C-cycle tended to increase in the presence of earthworms, when litter with high C-N ratio was provided as a food source. Especially in the miscanthus treatments, less N might remain for enzymatic degradation, indicating that N competition between earthworms and microorganisms may vary between different litter types. Especially, an expansion of miscanthus in agricultural management systems might result in a reduced microbial activity and a higher N deficit for microorganisms in soil.     

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.     

Production of lignocellulose-degrading enzymes and degradation of leaf litter by saprotrophic basidiomycetes isolated from a Quercus petraea forest

Due to the production of lignocellulose-degrading enzymes, saprotrophic basidiomycetes can significantly contribute to the turnover of soil organic matter. The production of lignin- and polysaccharide-degrading enzymes and changes of the chemical composition of litter were studied with three isolates from a Quercus petraea forest. These isolates were capable of fresh litter degradation and were identified as Gymnopus sp., Hypholoma fasciculare and Rhodocollybia butyracea. Within 12 weeks of incubation, H. fasciculare decomposed 23%, R. butyracea 32% and Gymnopus sp. 38% of the substrate dry mass. All fungi produced laccase and Mn-peroxidase (MnP) and none of them produced lignin peroxidase or other Mn-independent peroxidases. There was a clear distinction in the enzyme production pattern between R. butyracea or H. fasciculare compared to Gymnopus sp. The two former species caused the fastest mass loss during the initial phase of litter degradation, accompanied by the temporary production of laccase (and MnP in H. fasciculare) and also high production of hydrolytic enzymes that later decreased. In contrast, Gymnopus sp. showed a stable rate of litter mass loss over the whole incubation period with a later onset of ligninolytic enzyme production and a longer lasting production of both lignin and cellulose-degrading enzymes. The activity of endo-cleaving polysaccharide hydrolases in this fungus was relatively low but it produced the most cellobiose hydrolase. All fungi decreased the C/N ratio of the litter from 24 to 15-19 and Gymnopus sp. also caused a substantial decrease in the lignin content. Analytical pyrolysis mass spectrometry of litter decomposed by this fungus showed changes in the litter composition similar to those caused by white-rot fungi during wood decay. These changes were less pronounced in the case of H. fasciculare and R. butyracea. All fungi also changed the mean masses of humic acid and fulvic acid fractions isolated from degraded litter. The humic acid fraction after degradation by all three fungi contained more lignin and less carbohydrates. Compared to the decomposition by saprotrophic basidiomycetes, litter degradation in situ on the site of fungal isolation resulted in the relative enrichment of lignin and differences in lignin composition revealed by analytical pyrolysis. It can most probably be explained by the participation of non-basidiomycetous fungi and bacteria during natural litter decomposition.     

Scots pine litter decomposition along drainage succession and soil nutrient gradients in peatland forests, and the effects of inter-annual weather variation

Peatlands form a large carbon (C) pool but their C sink is labile and susceptible to changes in climate and land-use. Some pristine peatlands are forested, and others have the potential: the amount of arboreal vegetation is likely to increase if soil water levels are lowered as a consequence of climate change. On those sites tree litter dynamics may be crucial for the C balance. We studied the decomposition of Scots pine (Pinus sylvestris L.) needle and root litter in boreal peatland sites representing gradients in drainage succession (succession following water level drawdown caused by forest drainage) and soil nutrient level during several years of varying weather conditions. Neither gradient had an unambiguous effect on litter mass loss. Mass loss over 2 years was faster in undrained versus drained sites for both needle litter, incubated in the moss layer, and fine root litter, incubated in 0-10 cm peat layer, suggesting moisture stress in the surface layers of the drained sites limited decomposition. Differences among the drained sites were not consistent. Among years, mass loss correlated positively with precipitation variables, and mostly negatively or not at all with temperature sum. We concluded that a long-term water level drawdown in peatlands does not necessarily enhance decay of fresh organic matter. Instead, the drained site may turn into a ‘large hummock-system’ where several factors, including litter quality, relative moisture deficiency, higher acidity, lower substrate temperature, and in deeper layers also oxygen deficiency, may interact to constrain organic matter decomposition. Further, the decomposition rates may not vary systematically among sites of different soil nutrient levels following water level drawdown. Our results emphasize the importance of annual weather variations on decomposition rates, and demonstrate that single-period incubation studies incorporate an indeterminable amount of temporal variation.     

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.     

Influence of leaf litter types on microbial functions and nutrient status of soil: Ecological suitability of forest trees for afforestation in tropical laterite wastelands

The impact of forest tree leaf litters on microbial activity and nutrient status of red laterite soil was tested for the ecological suitability of Cassia siamea, Shorea robusta, Acacia auriculiformes and Dalbergia sissoo, which are typically used for afforestation of wastelands in eastern India. The objectives were to compare seasonal variation in soil enzyme activity in 30-years old afforested sites, and to study nutrient status and microbial biomass and function during short-term in-situ incubation of litter in decomposition pits. In afforested soils, enzyme activities significantly varied between litters and seasons. All enzyme activity except invertase dominated in the soils containing Dalbergia and Cassia litters compared to the others. The seasonal effect was enzyme-dependent, with amylase and cellulase reaching peaks during the rainy season but invertase activity showed a reverse trend with lowest values in rainy season, except in Acacia soil, and protease activity was lowest in the soil containing Cassia and Dalbergia during the rainy season. Dehydrogenase activity was negligible in the soils containing Shorea and Acacia, but remained high with respect to Dalbergia and Cassia during all seasons. The decomposition pit study showed significant increase of soil nutrients with respect to litter types and intervals, except with respect to electrical conductivity. Cassia and Dalbergia litters enabled notable increase of soil nutrients than Shorea and Acacia. The soil enzyme activity, in general, increased with duration of litter decay, but microbial biomass C (MBC) decreased over time except in Shorea. Therefore, the enzyme rates normalized to the MBC indicated inverse relations for all enzymes, except in the soil containing Shorea. A positive relationship existed between MBC and soil respiration in Cassia, Acacia and Dalbergia. Analysis of variance revealed main effects of litter types for increasing protease, MBC and CO₂ output, and a main effect of intervals for enhancing enzymes other than cellulase. Rates of soil respiration were greater in soils contain Cassia and Dalbergia, and showed significant differences between litters and between intervals. All enzymes were significantly correlated with electrical conductivity, organic carbon and available phosphorus contents, and all enzymes except invertase were correlated with nitrate concentrations. The acidic soil pH did not affect enzyme activities, and soil nutrients exerted only weak effect on MBC and respiration. Our study showed that leaf litters of Cassia and Dalbergia trees improved the nutrient status and microbial activity in soil more so than Shorea and Acacia litters, and therefore, afforestation using Cassia and Dalbergia trees may be particularly suitable for soil restoration in tropical laterite wastelands.     

The role of Armadillidium vulgare (Isopoda: Oniscidea) in litter decomposition and soil organic matter stabilization

We studied the effects of the terrestrial isopod Armadillidium vulgare on organic matter decomposition and stabilization in a long-term (65-week) laboratory experiment. We quantified the microbial activity in leaf litter (Acer pseudoplatanus) which did not come into contact with isopods, in A. vulgare feces produced from the same litter, and in unconsumed leftover of this litter. Freshly fallen leaf litter and up to 3 day old feces and leftover of litter were used. All materials were air dried immediately after collection and rewetted 1 day before use. Simultaneously, we measured how microbial activity in litter and feces are affected by fluctuations in humidity and temperature and by the addition of easily decomposed substances (starch and glucose).Microbial respiration was lower in feces than in litter or unconsumed leaf fragments. At the same time, moisture and temperature fluctuations and addition of glucose or starch increased respiration much more in litter than in feces. The results indicate that the processing of litter by A. vulgare reduces microbial respiration and reduces the sensitivity of microbial respiration to environmental fluctuations. ¹³C NMR spectra from feces indicated preferential loss of polysaccharide-carbon and accumulation of lignin with some modification to the aromatic-carbon. TMAH-Py-GC MS showed that lignin content was higher in feces than in litter and that lignin quality differed between the two substrates. Guaiacyl units were depleted in the feces, which indicated breakdown of guaiacyl associated with gut passage. As a conclusion, the results suggest that this common isopod greatly affects leaf litter decomposition. Decomposition of isopod feces in a long-term experiment is lower than litter decomposition which may support stabilization of organic matter in soil. This is caused mainly due to higher content of aromatic carbon in feces, which may cause its considerable resistance to bacterial degradation.     

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.     

Temperature and substrate controls on microbial phospholipid fatty acid composition during incubation of grassland soils contrasting in organic matter quality

Soil incubations are often used to investigate soil organic matter (SOM) decomposition and its response to increased temperature, but changes in the activity and community composition of the decomposers have rarely been included. As part of an integrated investigation into the responses of SOM components in laboratory incubations at elevated temperatures, fungal and bacterial phospholipid fatty acids (PLFAs) were measured in two grassland soils contrasting in SOM quality (i.e. SOM composition), and changes in the microbial biomass and community composition were monitored. Whilst easily-degradable SOM and necromass released from soil preparation may have fuelled microbial activity at the start of the incubation, the overall activity and biomass of soil microorganisms were relatively constant during the subsequent one-year soil incubation, as indicated by the abundance of soil PLFAs, microbial respiration rate (r), and metabolic quotient (qCO₂). PLFAs relating to fungi and Gram-negative bacteria declined relative to Gram-positive bacteria in soils incubated at higher temperatures, presumably due to their vulnerability to disturbance and substrate constraints induced by faster exhaustion of available nutrient sources at higher temperatures. A linear correlation was found between incubation temperatures and the microbial stress ratios of cyclopropane PLFA-to-monoenoic precursor (cy17:0/16:1ω7c and cy19:0/18:1ω7c) and monoenoic-to-saturated PLFAs (mono/sat), as a combined effect of temperature and temperature-induced substrate constraints. The microbial PLFA decay patterns and ratios suggest that SOM quality intimately controls microbial responses to global warming.     

Microbial community utilization of added carbon substrates in response to long-term carbon input manipulation

The chemical composition and quantity of plant inputs to soil are primary factors controlling the size and structure of the soil microbial community. Little is known about how changes in the composition of the soil microbial community affect decomposition rates and other ecosystem functions. This study examined the degradation of universally ¹³C-labeled glucose, glutamate, oxalate, and phenol in soil from an old-growth Douglas-fir (Pseudotsuga menziesii)—western hemlock (Tsuga heterophylla) forest in the Oregon Cascades that has experienced 7 y of chronic C input manipulation. The soils used in this experiment were part of a larger Detritus Input and Removal Treatment experiment and have received normal C inputs (control), doubled wood inputs, or root and litter input exclusion (no inputs). Soil from the doubled wood treatment had a higher fungal:bacterial ratio, and soil from the no inputs treatment had a lower fungal:bacterial ratio, than the control soil. Differences in the utilization of the compounds added to the field-manipulated soils were assessed by following the ¹³C tracer into microbial biomass and respiration. In addition, ¹³C-phospholipid fatty acids (PLFA) analysis was used to examine differential microbial utilization of the added substrates. Glucose and glutamate were metabolized similarly in soils of all three litter treatments. In contrast, the microbial community in the double wood soil respired more added phenol and oxalate, whereas microbes in the no inputs soil respired less added phenol and oxalate, than the control soil. Phenol was incorporated primarily into fungal PLFA, especially in soil of the double wood treatment. The addition of all four substrates led to enhanced degradation of soil organic matter (priming) in soils of all three litter treatments, and was greater following the addition of phenol and oxalate as compared to glucose and glutamate. Priming was greater in the no inputs soil as compared to the control or doubled wood soils. These results demonstrate that altering plant inputs to soil can lead to changes in microbial utilization of C compounds. It appears that many of these changes are the result of alteration in the size and composition of the microbial community.     

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.     

Effects of nitrogen availability on microbial activities,densities and functional diversities involved in the degradation of a Mediterranean evergreen oak litter (Quercus ilex L.)

The effect of available nitrogen N ((NH₄)₂S0₄) amendments on various microbial variables in three different layers (OhLn, OhLv, OhLf) of a Mediterranean litter profile under an evergreen oak forest (Quercus ilex L.) were studied. Since Mediterranean litters are generally N limiting, the goal of the study was to understand how low (0.1 and 1%) and high (5 and 10%) N amendments impact specific biological variables such as hyphal length, community-level-catabolic-profiles (CLCPs) in ECO and FF Biolog? plates, basal respiration, enzymatic activities (i.e. alkaline phosphatases (AlP), laccases, peroxidases and cellulases), and laccase and cellulase isoforms from three different litter layers. Results indicated that the effects of N amendments occurred over very short incubation time (3 d), and varied depending on N concentration and litter organic matter (OM) quality (i.e. depth). Thus, it appeared that the more active layer was the intermediate (OhLv) layer, which probably contained the most labile and available C pools. As a consequence, OhLv was also the layer showing globally the more intensive microbial responses following low N amendments. Indeed, in this layer, low N supplies caused several marked increases in enzymes activities (i.e. laccases, cellulases and alkaline phosphatases), hyphal length and isoenzyme patterns, suggesting a microbial reallocation of C to biomass and enzyme production. On the contrary, high N supplies resulted in adverse effects on almost all the variables, suggesting repression or cytotoxic phenomena.     

Decomposition of leaf litter from chitinase transgenic silver birch (Betula pendula) and effects on decomposer populations in a field trial

Although the area under cultivation of genetically modified plants (GMPs) has substantially increased during the last decade, the effects of transgenic organisms on ecosystem processes (such as litter decomposition and nutrient cycling) largely remain unknown. In this study, the decomposition of leaf litter from transgenic birch trees (Betula pendula) expressing sugar beet chitinase IV gene was studied in a field experiment. Eight chitinase transgenic lines and a non-transgenic control were included in the study. The decomposition of these litters was investigated by studying: (i) litter mass loss, (ii) fungal (litter ergosterol content) and total microbial biomass (SIR) and their activity (basal respiration), and (iii) the effects of transgenic litter on microbial-feeding soil fauna (number of nematodes and abundance of different functional groups). At the end of the study (8 and 11 months after establishment) mass loss of chitinase transgenic leaf litter did not differ from that of non-transgenic control trees. Similarly, no differences in either the fungal or total microbial biomass between the treatments were recorded. A single transgenic line showing high chitinase IV expression differed significantly from the controls in the mean number of nematodes. The nematode populations in this litter showed distinct temporal dynamics compared to the controls, thus indirectly indicating microbial differences in the litter. The results of this study indicate that conceivable changes, possibly derived from pleiotropic effects due to gene modification, in the litter quality due to gene transformation are either absent or too weak to affect the decomposability of the litter in the soil.     

Carbon dioxide production and oxygen consumption during the early decomposition of different litter types over a range of temperatures in soil‐inoculated quartz sand

Oat straw, hay, and alfalfa litter, differing in microbial colonization and recalcitrance, were added to organic matter–free quartz sand (5 mg C [g material]^–1) and incubated in the laboratory at 5°C, 10°C, 15°C, 20°C, and 25°C. Different incubation periods were chosen so that theoretically the same amounts of CO₂ would be produced and the same amounts of O₂ would be consumed for each litter type. It was investigated whether Q₁₀ values (change in respiration rate between two temperatures) increase with decreasing temperature and how much these Q₁₀ values and also the respiratory quotient (RQ: mol CO₂/mol O₂) depend on the litter type. The sums of CO₂‐C evolved and O₂ consumed, but also the contents of microbial biomass C and microbial biomass N showed a nearly 7‐fold increase in the order oat straw < hay < alfalfa litter. In contrast, the ratio of the fungal cell‐membrane component ergosterol to microbial biomass C was highest in the oat straw (4.1‰) and lowest in the alfalfa litter (0.2‰). This ratio reached a similar level between 5°C and 15°C (1.9‰), significantly higher (p = 0.01) than the level at 20°C (0.9‰). Respiration was similar between 20°C and 25°C, with a mean Q₁₀ value of 1.9. The use of temperature rate‐modifying factors suggested by the carbon‐turnover model ROTHC revealed that the incubation period for similar respiration rates was underestimated at 5°C and overestimated at 25°C. The lignin‐poor and protein‐rich alfalfa litter showed the highest Q₁₀ values of the three litter types in the medium temperature range of 10°C to 20°C. In contrast, the lignin‐rich and protein‐poor oat straw showed significantly highest Q₁₀ values at 5°C and 25°C in comparison with the other two litter types. The RQ was significantly highest in the hay litter (1.05) and in comparison with alfalfa litter (0.97) and oat straw (0.92). Strong temperature‐dependent variations in Q₁₀ values and respiratory quotients suggest interactions between litter quality, microbial colonization of litter, and temperature, which warrants further investigation.