Effect of N addition,moisture, and temperature on soil microbial respiration and microbial biomass in forest soil at different stages of litter decomposition

Purpose

The objective of the present study was to investigate the interactive effects of nitrogen (N) addition, temperature, and moisture on soil microbial respiration, microbial biomass, and metabolic quotient (qCO₂) at different decomposition stages of different tree leaf litters.

Materials and methods

A laboratory incubation experiment with and without litter addition was conducted for 80 days at two temperatures (15 and 25 °C), two wetting intensities (35 and 50 % water-filled porosity space (WFPS)) and two doses of N addition (0 and 4.5 g N m^?2, as NH₄NO₃). The tree leaf litters included three types of broadleaf litters, a needle litter, and a mixed litter of them. Soil microbial respiration, microbial biomass, and qCO₂ along with other soil properties were measured at two decomposition stages of tree leaf litters.

Results and discussion

The increase in soil cumulative carbon dioxide (CO₂) flux and microbial biomass during the incubation depended on types of tree leaf litters, N addition, and hydrothermal conditions. Soil microbial biomass carbon (C) and N and qCO₂ were significantly greater in all litter-amended than in non-amended soils. However, the difference in the qCO₂ became smaller during the late period of incubation, especially at 25 °C. The interactive effect of temperature with soil moisture and N addition was significant for affecting the cumulative litter-derived CO₂-C flux at the early and late stages of litter decomposition. Furthermore, the interactive effect of soil moisture and N addition was significant for affecting the cumulative CO₂ flux at the late stage of litter decomposition but not early in the experiment.

Conclusions

This present study indicated that the effects of addition of N and hydrothermal conditions on soil microbial respiration, qCO₂, and concentrations of labile C and N depended on types of tree leaf litters and the development of litter decomposition. The results highlight the importance of N availability and hydrothermal conditions in interactively regulating soil microbial respiration and microbial C utilization during litter decomposition under forest ecosystems.

     

Temperature and moisture dependence of decomposition rates of hardwood and coniferous leaf litter

Samples of fresh (autumn) and of year-old (late summer) deciduous forest-leaf litter and humus, and of Douglas fir fine litter and humus, were wetted to known moisture content, nominally between 200 and 40% water (dry basis), and maintained at constant temperatures between 10° and 40°C. Rates of CO₂ production were measured by KOH absorption and titration. Decomposition rate was found to be a linear function of log — (water potential), and to approach a maximum near 40°C. The temperature-dependence was consistent with models based on irreversible heat inactivation of a rate-controlling enzyme, also with Eyring's “absolute reaction rate” theory for reactions controlled by a reversibly inactivated enzyme. Activation energies were 66.8–67.3 kJ mol⁻¹ for litter, and 61.4–67.5 kJ mol⁻¹ for humus decomposition; for enzyme inactivation energies were 150–154 kJ mol⁻¹.     

Rainfall manipulation effects on litter decomposition and the microbial biomass of the forest floor

Simple structures aimed at regulating the amount of rain water dropping into the forest floor were installed to determine the impact of rainfall on leaf litter mass loss, respiration rates, microbial biomass C (C_mic) and metabolic quotient (qCO₂). The rainfall manipulation treatments were (I) fully covered (100% reduction); (II) partially covered (50% reduction) and (III) control (fully exposed). Using the litterbag technique, the mass losses of covered Quercus serrata, Quercus acutissima, Acer rufinerve and Pinus densiflora leaf litter were reduced (P<0.01) by 19–26% compared to fully exposed litter. A positive linear relationship (r=0.90; P<0.0001) between litter C_mic and mass loss was noted across all litter types and covering regimes. The mass losses in fully exposed litter were attributed to the leaching effect of rainfall coupled with the synergistic actions of microbes and soil fauna, as suggested by their respiration and microbial biomass. In the covered litter, C_mic was generally reduced (P<0.01) while fully and partially exposed litter were comparable (P>0.05). On the other hand, respiration rates and qCO₂ were variable and showed no consistent treatment effect except for respiration rates at 3 months. Similarly, soil respiration rates and C_mic were not consistently affected by cover treatments. Evidently, the zero-rainfall condition negatively affected some biological processes in the litter layer but sporadically affected soil processes. The absence of rainfall, even if the soil moisture content was maintained, could affect organic matter turnover in the forest floor.     

Soil characteristics determine soil carbon and nitrogen availability during leaf litter decomposition regardless of litter quality

Climate and litter quality have been identified as major drivers of litter decomposition, but our knowledge of how soil characteristics (e.g. microbial community and chemical properties) determine carbon (C) and nitrogen (N) availability derived from the decomposition of litter of different qualities is still scarce. We conducted a microcosm experiment to evaluate how soils with contrasting microbial communities and soil properties (denoted Soils A and B hereafter, where Soil B has higher bacterial and fungal abundance, fungal:bacterial ratio, and organic C than Soil A) determine the availability of soil C (carbohydrates, proteins, amino acids and phenols) and N (dissolved organic and inorganic N, microbial biomass N and available N) during the decomposition of litter of contrasting quality (C:N ratios ranging from 20 to 102). We also evaluated the relative importance of soil characteristics and litter quality as drivers of C and N inputs to the soil during this process. Overall, higher soil C and N availability after litter decomposition was found in Soil B than in Soil A. Soil characteristics had a higher positive effect on soil C and N contents than litter quality during litter decomposition. We also found that changes in N availability and organic matter quality registered after litter decomposition, linked to different soil characteristics, were able to promote dissimilarities in the potential mineralization rates. In conclusion, our study provides evidence that soil characteristics (e.g. microbial communities and chemical properties) can be more important than litter quality in determining soil C and equally important for N availability during the decomposition of leaf litter.     

Accumulation and solubility of metals during leaf litter decomposition in non-polluted and polluted soil

The decomposition of alder ( Alnus glutinosa ) and poplar ( Populus tremula ) leaf litter placed in direct contact with non-polluted and metal-polluted soil was investigated over 25 months in a controlled model-ecosystem experiment using the litterbag method. In addition to mass loss, we monitored the total and soluble concentrations of carbon, Cu, Zn, Cd and Pb. Leaves from trees grown on polluted soil had larger initial Zn, Cd and dissolved organic carbon concentrations. Neither the origin of the leaves (from trees grown on non-polluted or polluted soil) nor the placement of the leaves in polluted or unpolluted soil affected the decomposition process. Total metal contents increased in leaves placed on polluted soil over time. The solubility of metals in the leaf litter decreased over time, indicating that leaves acted as a temporary pool for metals from the soil in direct contact with the leaves. The sorbed metals were strongly bound in the litter even after two years of decomposition. The strong binding and thus reduced bioavailability of the metals provides an explanation for why they had no observable effects on litter decomposition.     

Differential effects of lichens and mosses on soil enzyme activity and litter decomposition

In the New Jersey Pinelands, canopy gaps in the pine-dominated forest support patches of lichens, mosses, and caespitose grasses. We tested the hypotheses that non-vascular plants and lichens can affect nutrient cycling processes and that mosses and lichens would differ from each other. We predicted that (1) lichen tissues would decompose more slowly than pine or moss tissues, (2) all plant materials would decompose more slowly beneath lichens than beneath mosses, and (3) soil enzyme activities would be higher under lichens than under mosses or grasses, reflecting greater nutrient limitation. We compared rates of decomposition of the litter of Pinus rigida and moss and lichen tissues, and measured soil enzyme activities responsible for nutrient mineralization from litter (acid and alkaline phosphatases, chitinase, β-glucosidase, aminopeptidase, and phenol oxidase) under three types of groundcover (lichens, mosses, and grasses) and unvegetated soil at two sites. While groundcover affected enzyme activities, the patterns of enzyme activities differed markedly between the two sites. In general, the enzyme activities were uniformly low. Decomposition rates were more strongly affected by the groundcover than by litter materials. While all litters tended to decompose more slowly under lichens than under mosses, supporting one of our initial hypotheses, the rates of decomposition were markedly different between the two sites. These results suggest that while mosses and lichens create patches of different soil function in both sites, the differences between the sites in unknown factors cause the enzyme activities and decomposition rates to differ.     

Interactive effects of warming, soil humidity and plant diversity on litter decomposition and microbial activity

Human activity has induced a multitude of global changes that are likely to affect the functioning of ecosystems. Although these changes act in concert, studies on interactive effects are scarce. Here, we conducted a laboratory microcosm experiment to explore the impacts of temperature (9, 12 and 15 °C), changes in soil humidity (moist, dry) and plant diversity (1, 4, 16 species) on soil microbial activity and litter decomposition.We found that changes in litter decomposition did not mirror impacts on microbial measures indicating that the duration of the experiment (22 weeks) may not have been sufficient to determine the full magnitude of global change effects. However and notably, changes in temperature, humidity and plant litter diversity/composition affected in a non-additive way the microbial parameters investigated. For instance, microbial metabolic efficiency increased with plant diversity in the high moisture treatment but remained unaffected in low moisture treatment suggesting that climate changes may mask beneficial effects of biodiversity on ecosystem functioning. Moreover, litter decomposition was unaffected by plant litter diversity/composition but increased with increasing temperature in the high moisture treatment, and decreased with increasing temperature in the low moisture treatment.We conclude that it is inevitable to perform complex experiments considering multiple global change agents in order to realistically predict future changes in ecosystem functioning. Non-additive interactions highlight the context-dependency of impacts of single global change agents.     

Soil moisture and soil-litter mixing effects on surface litter decomposition: A controlled environment assessment

Recent studies suggest the long-standing discrepancy between measured and modeled leaf litter decomposition in drylands is, in part, the result of a unique combination of abiotic drivers that include high soil surface temperature and radiant energy levels and soil-litter mixing. Temperature and radiant energy effects on litter decomposition have been widely documented. However, under field conditions in drylands where soil-litter mixing occurs and accelerates decomposition, the mechanisms involved with soil-litter mixing effects are ambiguous. Potential mechanisms may include some combination of enhanced microbial colonization of litter, physical abrasion of litter surfaces, and buffering of litter and its associated decomposers from high temperatures and low moisture conditions. Here, we tested how soil-litter mixing and soil moisture interact to influence rates of litter decomposition in a controlled environment. Foliar litter of two plant species (a grass [Eragrostis lehmanniana] and a shrub [Prosopis velutina]) was incubated for 32 weeks in a factorial combination of soil-litter mixing (none, light, and complete) and soil water content (2, 4, 12% water-filled porosity) treatments. Phospholipid fatty acids (PLFAs) were quantified one week into the experiment to evaluate initial microbial colonization. A complementary incubation experiment with simulated rainfall pulses tested the buffering effects of soil-litter mixing on decomposition.Under the laboratory conditions of our experiments, the influence of soil-litter mixing was minimal and primarily confined to changes in PLFAs during the initial stages of decomposition in the constant soil moisture experiment and the oscillating soil moisture conditions of the rainfall pulse experiment. Soil-litter mixing effects on CO₂ production, total phospholipid concentrations, and bacterial to total PLFA ratios were observed within the first week, but responses were fairly weak and varied with litter type and soil moisture treatment. Across the entire 32-week incubation experiment, soil moisture had a significant positive effect on mass loss, but soil-litter mixing did not. The lack of strong soil-litter mixing effects on decomposition under the moderate and relatively constant environmental conditions of this study is in contrast to results from field studies and suggests the importance of soil-litter mixing may be magnified when the fluctuations and extremes in temperature, radiant energy and moisture regimes common dryland field settings are in play.     

Mass loss measurements and statistical models to predict decomposition of leaf litter in a boreal aspen forest

Abstract

In a southern boreal aspen forest located in Saskatchewan, Canada, we examined decomposition rates of leaf litter from trembling aspen (Populus tremuloides Michx.), hazel (Corylus cornuta March.), and a mixture of different species over a six‐month period. Mass loss was measured in the field using the litter bag method. The greatest mass losses occurred during the first month regardless of litter type. On average, mass loss during the first 28 days was 3.2 g#lbkg^‐1#lbd^‐1 for the aspen leaves, 4.4 for hazel leaves and 4.9 for the mixture. The initial rapid loss of weight is attributed to leaching and decomposition of water soluble material. The decomposition rates of the leaf litter were related to water‐soluble organic carbon and nitrogen content, and C:N ratio. Several models were used to describe mass loss of the aspen, hazel, and mixed leaf litter at the early stages of decomposition. A single model was not found to be appropriate to describe decomposition of all leaf‐litter types. A second order model provided the best fit for the aspen litter decomposition, while the logarithmic model best described the decomposition of hazel and mixed leaf litter.     

Dominant effects of litter substrate quality on the difference between leaf and root decomposition process above- and belowground

Initial decomposition rates, changes in organic chemical components (acid-insoluble fraction, holocellulose, polyphenols, soluble carbohydrates) and nutrient dynamics (K, Mg, Ca, P, N) were examined for fine roots and leaves of Japanese cypress (Chamaecyparis obtusa). Litterbag experiments designed to evaluate the relative effects of litter type and position of litter supply in the soil were carried out, considering that root and leaf litter typically occupy different locations and have different substrate qualities. Litterbags of roots and leaves were placed at two positions (on the soil surface and in the humus layer), and collected every 3 months over one year. The mass loss rate and N release were slower during root decomposition in the humus layer than during leaf decomposition on the soil surface. These differences between root and leaf decomposition were mainly caused by the litter type, and the effect of the position on decomposition was relatively small. Root litter was less influenced by position related effects, such as differences in humidity, than leaf litter, and this recalcitrant trait to environmental effects may be responsible for the slower mass loss rate and N release in root decomposition. The results of the present study suggest that fine roots are persistent in the soil and serve an important role in N retention in forest ecosystems because of their litter substrate quality.     

温度与土壤水分对有机碳分解速率的影响

试验研究温度与土壤水分对有机碳分解速率的影响,结果表明:相同水分条件下,培育初期(1～20d)各处理CO2排放速率较高,相对值表现为35℃处理>25℃处理>15℃处理>5℃处理;随培育时间延长(>20d),CO2排放速率渐趋平稳。相同温度条件下,30%～90%田间持水量时培育初期(1～20d)各处理CO2排放速率初始值较高,之后降低,当趋于某一定值时,相对值大小随土壤水分含量增加而增加。相同温度与土壤水分条件下,CO2排放速率相对值大小随土壤有机碳含量增加而增加。     

Determinants of leaf litter patchiness in mixed species New Jersey pine barrens forest and its possible influence on soil and soil biota

 We have identified the importance of ground layer ericaceous shrub density as a determinant of leaf litter patch size in upland oak/pine communities of the New Jersey pine barrens. Litter patch area is directly proportional to the number of ericaceous stems. This observation has been confirmed by experimentation where leaf litter patches accumulated under artificial stems for a period of 2 years. Leaf litter patches of different sizes contain differing proportions of leaf species. Large patches contain a significantly higher proportion of oak leaves than small patches. Difference in physical structure of large and small patches, due to leaf species composition and due to differential leaf chemistries, result in differences in soil characteristics and soil biota under the patches. Soil moisture and organic matter content of upper soil layers are greater under large litter patches than under small ones. We have preliminary evidence that these changes influence the community structure of ectomycorrhizae developing under patches of different sizes. Received: 12 April 1999     

Effect of litter quality on its decomposition in broadleaf and coniferous forest

Recently there has been much interest in the effect of litter mixing as well as the effect of different forest habitats on the decomposition process. Our aim was to test two hypotheses: high quality litter promotes decomposition of poor quality litter, and litter decomposes faster in broadleaf than in coniferous forest. We conducted a litter mixing experiment using litterbags placed in two forest floors, in which treatments consisted of litter monocultures of each of two campy species (Castanopsis eyrei and Pinus massoniana), as well as mixtures of these two species. The results showed that C. eyrei leaves decomposed significantly faster in the coniferous habitat than in their native habitat. On the other hand, P. massoniana needles decomposed significantly faster in their native coniferous habitat than in the broadleaf habitat. In our experiment we found that the mixture had different effect on different quality litter. P. massoniana needles (poor quality) had a positive effect on the decomposition of C. eyrei leaves (high quality), while C. eyrei leaves had a negative effect on the decomposition of P. massoniana needles in the mixture case in both broadleaf and coniferous habitats. The diversity of the fungi identified from different litters varied among treatments and the mass loss was positively correlated with the Shannon–Weaver diversity index of fungi. It is suggested that fungi may be one of the major drivers to control the decomposition process.     

Effects of the ecological restoration practices of prescribed burning and mechanical thinning on soil microbial enzyme activities and leaf litter decomposition

The effects of ecological restoration on belowground processes such as decomposition are generally unknown. To assess the immediate effects of prescribed fire and mechanical thinning on belowground processes, we measured the activities of five extracellular enzymes (phosphatase, β-glucosidase, β-N-acetylglucosaminidase, phenol oxidase, and lignin-peroxidase) in soils and on decomposing Quercus falcata leaf litter in unburned, burned, and burned and thinned plots in a mesic forest in northern Mississippi. Decomposition rates of Q. falcata leaf litter were also assessed at each plot. Soil phosphatase activity decreased after a prescribed burn and was related to an increase in soil organic matter in plots that had been burned. Soil β-N-acetylglucosaminidase activity increased after a burn, and was related to a decrease in leaf litter. Leaf litter enzyme activity showed no consistent patterns amongst treatments, or between individual enzymes, while decomposition rates of leaf litter were slightly accelerated in the treatment plots, but not significantly so. Decomposition rates were related to cumulative enzyme activity, with phenol oxidase and lignin-peroxidase having the highest apparent efficiencies in degrading the leaf material. Overall, the microbial degradation of Q. falcata leaf litter was more efficient in plots that were burned and thinned than in the other plots. Increases in the efficiency of litter decomposition coupled with reductions in litter inputs due to canopy thinning likely allows for increased solar penetration to the soil, and could promote the restoration of the shade-intolerant species that once dominated the understory. Post-burn increases in β-N-acetylglucosaminidase activity and decreases in phosphatase activity also suggest a potential shift in the soil community from phosphorus limitation to nitrogen limitation following a fire.     

Extractability of cellulases in forest litter and soil

Summary Cellulases in forest litter and soil occur in both bound and extractable forms. The proportion of total bound endocellulase activity (not extractable) increases during decomposition, whereas the proportion of bound exocellulase activity remains fairly constant. The proportions of bound enzymes differ among litter types with different chemical compositions. The proportion of bound activity is higher in mineral soil than in litter. We also investigated the effects of anion type (NaCl versus Na₂SO₄), concentration and pH on the extractability of cellulases and protein in two horizons of two forest soils. The extractability of cellulases increases as pH increases from 3.5 to 5.6. Anion type and concentration did not have consistent effects on extractability. However, there was a trend for higher extractability by sulfate than by chloride and with increasing salinity.     

Direct effects of litter decomposition on soil dissolved organic carbon and nitrogen in a tropical rainforest

To clarify how litter decomposition processes affect soil dissolved organic carbon (DOC) and soil dissolved nitrogen (DN) dynamics, we conducted a field experiment on leaf litter and collected DOC and DN from the underlying soil in a tropical rainforest in Xishuangbanna, southwest China. Principal components analysis (PCA) showed the first PCA axis (corresponding to degraded litter quantity and quality) explained 61.3% and 71.2% of variation in DOC and DN concentrations, respectively. Stepwise linear regression analysis indicated that litter carbon mass controlled DOC and hemicellulose mass controlled DN concentrations. Litter decomposition was the predominant factor controlling surface-soil DOC and DN dynamics in this tropical rainforest.     

Chemical and bacterial changes in a forest soil percolated with some amino acids and leaf litter extract

Chemical and bacterial changes in the forest soil percolated with glycine and glutamate solution and leaf litter extract were studied.

When the forest soil was percolated with 25 mM glycine or glutamate, it took about 20 days for the number of bacteria to reach the maximum in number and the amino acids to be completely ellhausted, the rate being much slower than that in the cultivated soil. Supplementation of 0.01% yeast extract to the percolate much rnhanced the growth of bacteria and degradation of glycine. In this percolation the metabolic pattern of glycine was comparable to that in the cultivated soil except for the absence of an appreciable amount of nitrate formation.

Growth of bacteria in the forest soil was also rapid when percolated with leaf litter extract and only 3 days were sufficient for reaching the maximum number. Neither ammonia nor nitrate was detected throughout the percolation period and only change observed in the solution was a slight rise in pH.

Characteristics of bacteria enriched in the forest soil percolated will) glycine with and without yeast extract, and leaf litter extract were studied, No one group was dominant in the soil before percolation. Bacteria enriched by glycine were almost occupied with Gram-negative nonmotile rods with rather complex nutritional requirement regarded as Achromobacter, which were characteristically unable to utilize glycine as a sole nitrogen source. When the soil was percolated with glycine supplemented with yeast extract, enriched bacteria were composed of many kinds. The soil percolated with leaf litter extract was occupied with bacteria with simple nutritional requirement, which were regarded as Pseudamonas.     

Ant-mediated effects on spruce litter decomposition, solution chemistry, and microbial activity

Forest management practices often generate clear-cut patches, which may be colonized by ants not present in the same densities in mature forests. In addition to the associated changes in abiotic conditions ants can initiate processes, which do not occur in old-growth stands. Here, we analyse the effects of ants and aphid honeydew on litter solution of Norway spruce, microbial enzyme activities, and needle decomposition in a field and greenhouse experiment during summer 2003. In the field, low ant densities had relatively little effects on litter solution 30 cm away from a tree trunk, but significantly increased organic carbon concentrations and decreased inorganic nitrogen concentrations next to a trunk where ants tend to build their nests. In a greenhouse experiment, the addition of ants to lysimeters containing spruce litter significantly increased dissolved organic carbon (DOC), dissolved organic nitrogen (DON), NH₄-N, NO₃-N and K concentrations in litter solutions compared to the control treatment, while the simulation of aphid infestation (addition of honeydew) significantly increased DOC as a direct result of honeydew leaching, and decreased inorganic N concentrations in leachates. The presence of ants resulted in a changed composition of dissolved organic matter (DOM) with more aromatic and complex compounds, and microbial enzyme activity was significantly higher in litter extracts from the ant treatment compared to the honeydew and control treatment. However, mass loss, litter %C and %N were not affected by ants or honeydew. Our results suggest that ants have a distinct and immediate effect on solution composition and microbial activity in the litter layer indicating accelerated litter decay whereas the effect of honeydew was insignificant.     

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.