Modeling of the thermal influence of fires on the physicochemical properties and microbial activity of litter in cryogenic soils

Periodic surface fires in the cryolithozone (the northern taiga subzone) are the main factor determining the qualitative and quantitative characteristics of the soil organic matter. The specific features of the changes in the physicochemical parameters and microbial activity of the organic horizons in the cryogenic soils under larch forests of the northern taiga after the impact of high temperatures were revealed. The temperatures of fires of different intensity were simulated in laboratory conditions. The thermal impact on the litter organic matter during the surface fires may increase the CO₂ emission from the surface of the soil in the postfire communities due to the destruction of organic compounds only for a short time. After fires of high intensity with strong mineralization of the litters, during a period of more than 1 month, the pyrogenic effect on the organic horizons of the soils under the larch forests of the cryolithozone determined the reduction of the CO₂ emissions in the freshly burned areas as compared to the intact stands.     

Transformation of microbial cenoses in soils of light coniferous forests caused by cuttings and fires in the Lower Angara River basin

The influence of surface fires and cutting on the quantitative and functional parameters of microbial cenoses in the soils of light coniferous forests in the Lower Angara River basin was studied. In the litters of soddy-podzolic soils under pine forests, the microbial biomass was 4080–4700 μg C/g; the basal respiration was 17.00–20.32 μg C-CO₂/g/h; and the qCO₂, 4.17–4.33 μg C-CO₂/mg C_mic/h. In the humus-accumulative horizon, these values were 880–1160 μg C/g, 2.48–4.12 μg C-CO₂/g/h, and 2.83–3.55 C-CO₂/mg C_mic/h, respectively. In the litter of the one-year-old felled area, the content of microbial biomass carbon was by two times lower; in the litter of burned plots, it was by 60–70% lower than in the litter of the control area. The intensity of the microbial respiration did not change proportionally to the microbial biomass content, which resulted in an imbalance between the processes of the organic matter mineralization-immobilization towards a release of CO₂ as evidenced by the increase of the qCO₂ values by 2–4 times. In the five-year-old felled area, at the stage of restoring the herbaceous vegetation, a tendency towards the stabilization of the destructive microbiological processes was revealed. In the felled areas, the high number of heterotrophic microorganisms, the reduced oligotrophy of the soil organic horizons, and the more intense microbiological mineralization of the organic matter were observed. The surface fires in the felled areas and forests significantly affected the structure and the number of ecological-trophic groups of microorganisms in the litters, the humus-accumulative horizons, and in the upper mineral soil layers. The maximal structural and functional disturbance in the soil microbial complex was found in the logged areas affected by fires.     

Influence of soil properties on the adsorption and mobility of metamitron

Abstract

The adsorption and mobility of herbicide, metamitron, in 41 soils samples from the province of Salamanca (Spain) was studied. Thirty‐four of the samples assayed were from irrigated soils and seven were from natural, uncultivated soils with organic matter contents above 3%. The correlations between the Freundlich K constants, Kd distribution coefficients, and Rf mobility parameters and the soils parameters were determined. Considering all the soils, the soils with organic matter contents above 2% or the soils with organic matter contents below 2%, significant correlations (p<0.001 to p<0.05) were found between K and Kd and the organic matter content of the soils. There were also a significant correlations (p<0.05) of K and Kd with clay+silt and clay contents of soils with organic matter contents below 2%. The adsorption of the herbicide by isolated soil components confirmed the results obtained with the soils and point to the importance of the exchangeable cation nature of the samples in the adsorption process. Based on Rf values obtained by TLC, the herbicide was found to be moderately mobile in 74% and mobile in 26% of the soils studied. The results of metamitron leaching by thin layer chromatography (TLC) and in undisturbed soil columns indicated the influence of organic matter content and of soil texture on the mobility of this herbicide.     

Pyrogenic changes in iron-illuvial podzols in the middle taiga of the Komi Republic

The changes in the morphological, physicochemical, and chemical properties of iron-illuvial podzols under middle-taiga dwarf shrub-green moss pine forests in the first months after fires are examined. It is shown that forest fires are accompanied by changes in the morphology of the litter horizons, their compaction, and by changes in the chemical properties of the soils. The data on the polycyclic aromatic hydrocarbons (PAHs) indicate a significant increase in the contents of chrysene, fluorene, naphthalene, pyrene, and anthracene in the Opyr(L) horizon in comparison with the litters in the background pine forest. The total PAH content in the O(F)pyr horizon increases mainly at the expense of di- and tri-nuclear PAHs (naphthalene and fluorene). The mineral soil horizons in the burnt area become enriched in the most mobile amphiphilic fractions of the organic matter, which is seen from the increase in the absolute and relative contents of the hydrophilic fractions, which might be represented by the products of combustion of fresh plant remains and litter.     

Changes in soil organic matter and net nitrogen mineralization in heathland soils,after removal,addition or replacement of litter from Erica tetralix or Molinia caerulea

Summary The effects of different litter input rates and of different types of litter on soil organic matter accumulation and net N mineralization were investigated in plant communities dominated by Erica tetralix L. or Molinia caerulea (L.) Moench. Plots in which the litter on the soil had repeatedly been removed were compared with plots in the same plant community in which litter had been added to the soil. In another treatment, litter was removed and replaced by litter from the other plant community. Net N mineralization was measured in situ after 5 years. Less soil organic matter and soil N was found in plots in which litter had been removed, compared with control plots, or plots to which litter had been added, but these differences were significant for the Erica sp. soils only. Plots in which litter had been replaced and control plots did not differ significantly in the amount of soil organic matter. However, in both plant communities, the differences agreed with the faster decomposition rate of Molinia sp. litter compared with Erica sp. litter. The gravimetric soil moisture content was correlated positively with the amount of soil organic matter, both in the Erica sp. soils and the Molinia sp. soils. Net N mineralization rates (g N m^-2) differed significantly between treatments for Erica sp. soils but no for Molinia sp. soils. For Erica sp. soils, net N mineralization rates increased with increasing amounts of soil organic matter and soil N. Replacing the litter with Molinia sp. litter (which differs in chemical composition) had no clear additional effect on the net N mineralization rate.     

Polycyclic aromatic hydrocarbons in pyrogenic soils of swampy landscapes of the Meshchera lowland

The composition and distribution of polycyclic aromatic hydrocarbons (PAHs) were studied in organomineral and organic soils of the Meshchera National Park. It was found that the background oligotrophic peat soils unaffected by fires in central parts of the bogs are characterized by the increased PAH concentrations due to their high sorption capacity. The fires of 2007 and 2010 resulted in the transformation of the plant cover and soil morphology, the formation of new horizons, and the change in the PAHs content and composition. Significant burn-off of organic matter was found in oligotrophic-eutrophic soils and resulted in the decrease of PAHs content after fire. Only partial burn-off of organic horizons and intense formation of PAHs were recorded in the soil with initially great thickness of peat horizons. Pyrogenic accumulation of PAHs was identified in organomineral soils of the marginal parts of bogs and of forest sites.     

Soils of postpyrogenic larch stands in Central Siberia: Morphology,physicochemical properties,and specificity of soil organic matter

Morphological features, physicochemical properties, and specific characteristics of the organic matter of cryozems (Cryosols) under postpyrogenic larch forests affected by fires 2, 6, 22, 55, and 116 years ago are considered. The morphological changes in the soils affected by fires are manifested by the burning of the upper organic horizons with preservation of pyrogenic features in the soils for more than a century after the fire. In the first years (2 and 6 years) after the fire, the acidity of the organic horizons and their base saturation become lower. The postpyrogenic soils are characterized by the smaller contribution of the organic horizons to the total pools of soil organic carbon. In the studied cryozems, the organic carbon content is correlated with the contents of oxalate-extractable iron and aluminum. A decrease in the content of water-soluble organic compounds in the soils is observed after the fires; gradually, their content increases upon restoration of the ground cover.     

Accumulation and transformation of organic matter in different-aged dumps from sand quarries

The accumulation and transformation of organic matter were studied in chronoseries of different-aged (3-, 10-, 20-, 30-, 43-, and 60-year-old) soils and a background (reference) plot. The ecogenetic succession of plants on sand quarry dumps was characterized. It was shown that the pedogenesis rate was closely related to the rate of phytocenosis development, and the thicknesses of organic and mineral horizons increased synchronously. The profile distribution of organic matter in young soils was estimated as an ectomorphic distribution, and the humus reserves in the mineral horizons of the same soils were comparable with the reserves of organic matter in the litters. The illuvial horizons of the soils under study played a significant role in the deposition of carbon dioxide; the resistance of organic matter to mineralization increased with age. In the soil chronoseries, the combustion heat of litter organic matter increased, as well as the content of energy accumulated in the litters. The composition of humus differed strongly between the eluvial and illuvial horizons; in the chronosequence, the relative content of humic acids increased in the E horizon, and that of fulvic acids increased in the B horizon. The effect of the phytocenosis on the soil was increasingly mediated with time. The accumulation and transformation of organic matter were the leading pedogenic processes at all stages.     

Pyrogenic transformation of organic matter in soils of forest bogs

Natural fires on forest bogs significantly affect all the groups and fractions of peat organic matter. The type and intensity of the fires are responsible for the depth of the pyrogenic transformation of peat. In the course of thermal destruction of peat organic matter, humus substances (humic acids in particular) are accumulated, which leads to changes in the type of humus; the humus reserves may increase by 1.5–8 times. Several ways of the formation of humus components related to the intensity of a fire are suggested. The regressive evolution of bog ecosystems caused by fires is a reversible process. The humus status of pyrogenically transformed horizons and their morphology are preserved within the peat deposit as a relic characteristic of the discrete metamorphosis of the soils.     

Tillage and amendment effects on soil carbon and nitrogen mineralization and phosphorus release

The influence of tillage and nutrient amendment management on nutrient cycling processes in soil have substantial implications for environmentally sound practices regarding their use. The effects of 2 years of tillage and soil amendment regimes on the concentrations of soil organic matter variables (carbon (C), nitrogen (N) and phosphorus (P)) and C and N mineralization and P release were determined for a Dothan fine-sandy loam soil in southeastern Alabama. Tillage systems investigated were strip (or conservation) and conventional tillage with various soil nutrient amendments that included no amendment, mineral fertilizer, and poultry waste (broiler litter). Surface soil (0–10 cm depth increment) organic matter variables were determined for all tillage/amendment combinations. Carbon and N mineralization and P release were determined on surface soils for each field treatment combination in a long-term laboratory incubation. Soil organic P concentration was 60% greater in soils that had been conventionally tilled, as compared with strip-tilled, both prior to and following laboratory incubation. Carbon and N mineralization results reflected the effects of prior tillage amendment regime, where soils maintained under strip-till/broiler litter mineralized the greatest amount of C and N. Determination of relative N mineralization indicated that strip tillage had promoted a more readily mineralizable pool of N (6.1%) than with conventional till (4.2%); broiler litter amendments had a larger labile N fraction (6.7%) than was found in soils receiving either mineral fertilizer (4.1%) or no amendment (4.7%). Tillage also affected P release measured during the incubation study, where approximately 20% more inorganic P was released from strip-tilled soils than from those maintained under conventional tillage. Greater P release was observed for amended soils as compared with soils where no amendment was applied. Results from this study indicate that relatively short-term tillage and amendment management can significantly impact C, N, and P transformations and transfers within soil organic matter of a southeastern US soil.     

Retention and extractability of phenol,cresol, and dichlorophenol exposed to two surface soils in the presence of horseradish peroxidase enzyme

The retention of phenol, o-cresol, 2,4-dichlorophenol (DCP), and their peroxidase-catalyzed polymerization products was evaluated on two surface soils. The extractability of the parent solutes and their polymerization products was also investigated. (14)C-Labeled radioisotopes were used to quantify the contaminant retained on soil as water-extractable, methanol-extractable, humic/fulvic (HA/FA) acid-bound, and soil/humin bound. Between 2 and 20% of the solute retained on soil after a 7-day contact period remained bound to the HA/FA and soil/humin components in unamended soils; in the presence of peroxidase this amount was as high as 40-75%. The alkali-extractable HA/FA component contained the largest fraction of radioactivity in peroxidase-amended soils. Whereas the soil organic matter content was the predominant factor controlling the extent of sorption of the parent phenols, the clay content and particle surface area appeared to contribute to the retention of the polymerization products. High molecular weight oligomers produced during peroxidase-mediated polymerization of phenols associate strongly with soil components and are likely incorporated into the soil organic matter via oxidative coupling reactions.     

Effects of addition of maize litter and earthworms on C mineralization and aggregate formation in single and mixed soils differing in soil organic carbon and clay content

C mineralization and aggregate stability directly depend upon organic matter and clay content, and both processes are influenced by the activity of microorganisms and soil fauna. However, quantitative data are scarce. To achieve a gradient in C and clay content, a topsoil was mixed with a subsoil. Single soils and the soil mixture were amended with 1.0 mg maize litter C g soil⁻¹ with and without endogeic earthworms (Aporrectodea caliginosa). The differently treated soils were incubated for 49 days at 15 °C and 40% water holding capacity. Cumulative C mineralization, microbial biomass, ergosterol content and aggregate fractions were investigated and litter derived C in bulk soil and aggregates were determined using isotope analyses. Results from the soil mixture were compared with the calculated mean values of the two single soils. Mixing of soil horizons differing in carbon and clay content stimulated C mineralization of added maize residues as well as of soil organic matter. Mixing also increased contents of macro-aggregate C and decreased contents of micro-aggregate C. Although A. caliginosa had a stimulating effect on C mineralization in all soils, decomposition of added litter by A. caliginosa was higher in the subsoil, whereas A. caliginosa decreased litter decomposition in the soil mixture and the topsoil. Litter derived C in macro-aggregates was higher with A. caliginosa than with litter only. In the C poor subsoil amended with litter, A. caliginosa stimulated the microbial community as indicated by the increase in microbial biomass. Furthermore, the decrease of ergosterol in the earthworm treated soils showed the influence of A. caliginosa on the microbial community, by reducing saprotrophic fungi. Overall, our data suggest both a decrease of saprotrophic fungi by selective grazing, burrowing and casting activity as well as a stimulation of the microbial community by A. caliginosa.     

NaOH‐extractable organic matter of andic soils from Galicia (NW Spain) under different land use regimes: a pyrolysis GC/MS study

The objective of this study was to determine to what extent the attenuation or loss of andic soil properties caused by land use change – from forest (FOR, average C content 118.2 ± 23.7 g kg^?1) to agricultural land (AGR, average C content 55.7 ± 16.7 g kg^?1) use – is reflected in soil organic matter (SOM) at the molecular level. For this, NaOH‐extractable SOM of A horizons from 17 soils developed on amphibolitic parent material in NW Spain was studied by pyrolysis gas chromatography spectrometry (Py‐GC/MS). We also included two buried andic A horizons (PAL, 2200 cal yr BP in age) on the same parent material, as a reference for the molecular composition of SOM from soils without recent litter additions. Organic matter of PAL soils had a composition largely different from that of superficial soils (FOR and AGR), with an important relative contribution of microbial polysaccharides and N‐compounds, and an absence of compounds that characterize fresh plant litter (e.g. lignins). In the superficial soils, the relative contribution of lignin‐derived compounds was greater in AGR than in FOR soils. Differences were also observed in the relative contribution of aliphatic compounds, FOR soils being enriched in this type of components compared with AGR soils. The results indicated that land use change from FOR to AGR, which was accompanied by a decrease in total SOM, resulted in an enrichment in primary SOM. The smaller relative abundance of primary SOM derivatives in andic FOR soils indicates that these compounds were quickly degraded in Andisols.     

Sorption of s-Triazine Herbicides in Organic Matter Amended Soils: Fresh and Incubated Systems

Fresh amendment of soil with sewage sludge and composted sewage sludge resulted in increased sorption of three s-triazine herbicides: atrazine, ametryn and terbuthylazine. The extent of increased sorption (as evaluated by sorption coefficients K_d or K_f) was a function of soil type, such that sorption in amended organic carbon-poor soil (0.4% OC) was more enhanced than in amended organic carbon-rich soil (1.55% OC). Despite significant differences between the organic amendments in terms of humic and fulvic acid content, humin content, soluble organic matter content, total organic matter content, and H/C and O/C atomic ratios, organic matter composition had no discernible effect on either sorption distribution coefficients or on isotherm linearity in amended soils. Soils amended with composted sludge had the same sorption potential as did soils amended with the analogous uncomposted sludge. After incubating soil-sludge mixtures for a year at room temperature, organic matter content decreased to original pre-amendment levels. Sorption coefficients for the three compounds similarly decreased to initial pre-amendment values. Organic carbon normalized sorption coefficients (K_oc) were essentially identical in the soils, amended soils, and incubated amended soils, indicating that sludge and compost derived organic matter does not have a significantly different sorption capacity as compared with the original soils, despite compositional differences.     

Soils of Low-Mountain Landscapes of North Karelia

Soils of low-mountain landscapes in the northwest of Karelia have been studied. The soil cover of the studied area is mainly represented by Al–Fe-humus soils (Podzols); thin soils (Leptosols) are widespread. Characteristic morphological features of all the studied soils are relatively shallow profiles, high stone content, and underlying by hard bedrock with fine earth material in crevices between large boulders. The studied soils have the high carbon and low nitrogen content, which points to unfavorable conditions of organic matter transformation. The content of most macro- and microelements is not high, which is typical for soils of the region; the content of copper and zinc exceeds the regional background two–three times. Regularities of the vertical zonality in the properties of soils of mountain ecosystems manifest themselves in decreasing thickness of the soil profile at higher altitude above sea level in parallel to decreasing thickness of the layer of loose rocks, while the stone content increases. In soils of the forest-tundra zone, the organomineral horizon with the high organic matter content is formed immediately under the forest litter. The litter horizon is the soils of this zone is characterized by increased concentrations of calcium, magnesium, phosphorus, and zinc.     

Use of pyrolysis/GC–MS combined with thermal analysis to monitor C and N changes in soil organic matter from a Mediterranean fire affected forest

In Mediterranean ecosystems, forest fires are a common phenomenon. They involve the transformation of vegetation and litter, leaving charred residues and so influencing the carbon cycle by changing (a) the amounts of soil organic matter and (b) the proportions within it of pools with differing stability. In addition to affecting C cycles, fires also affect the amounts of N within soil organic matter, and its availability.     

The biochemical transformation of oak (Quercus robur) leaf litter consumed by the pill millipede (Glomeris marginata)

Soil macrofauna play an essential role in the initial comminution and degradation of organic matter entering the soil environment and yet the chemical effects of digestion on leaf litter are poorly understood at the molecular level. This study was undertaken to assess the selective chemical transformations that saprophagous soil invertebrates mediate in consumed leaf litter. A number of pill millipedes (Glomeris marginata) were fed oak leaves (Quercus robur) after which the biomolecular compositions (lipids and macromolecular components) of the leaves and millipede faeces were compared using a series of wet chemical techniques and subsequent analysis by gas chromatography (GC) and gas chromatography-mass spectrometry (GC/MS). It was found that the concentrations of short chain (<C₂₀) n-alkanoic acids, sterols and triacylglycerols reduced dramatically in the millipede faeces relative to the leaf litter. Hydrolysable carbohydrates and proteins both decreased in concentration in the faeces, whereas similar yields of phenolic components were observed for the cupric oxidation products of lignin, although the oxygenated functionalities were affected by passage through the millipede gut, yielding a more highly condensed state for lignin. This shows that the chemical composition of fresh organic matter entering the soil is directly controlled by invertebrates feeding upon the leaf litter and as such that they are key contributors to the early stages of diagenesis in terrestrial soils.     

Adsorption and Desorption of Copper in Pasture Soils

Abstract

Copper (Cu) is bound strongly to organic matter, oxides of iron (Fe) and manganese (Mn), and clay minerals in soils. To investigate the relative contribution of different soil components in the sorption of Cu, sorption was measured after the removal of various other soil components; organic matter and aluminum (Al) and Fe oxides are important in Cu adsorption. Both adsorption and desorption of Cu at various pH values were also measured by using diverse pasture soils. The differences in the sorption of Cu between the soils are attributed to the differences in the chemical characteristics of the soils. Copper sorption, as measured by the Freundlich equation sorption constants [potassium (K) and nitrogen (N)], was strongly correlated with soil properties, such as silt content, organic carbon, and soil pH. The relative importance of organic matter and oxides on Cu adsorption decreased and increased, respectively, with increasing solution Cu concentrations. In all soils, Cu sorption increased with increasing pH, but the solution Cu concentration decreased with increasing soil pH. The cumulative amounts of native and added soil Cu desorbed from two contrasting soils (Manawatu and Ngamoka) during desorption periods showed that the differences in the desorbability of Cu were a result of differences in the physico‐chemical properties of the soil matrix. This finding suggests that soil organic matter complexes of Cu added through fertilizer, resulted in decreased desorption. The proportions of added Cu desorbed during 10 desorption periods were low, ranging from 2.5% in the 24‐h to 6% in the 2‐h desorption periods. The desorption of Cu decreased with increasing soil pH. The irreversible retention of Cu might be the result of complex formation with Cu at high pH.     

A new approach to calculate the particle density of soils considering properties of the soil organic matter and the mineral matrix

The particle density of soil (ρ_S) represents one of the soil's basic physical properties and it depends on the composition of both the mineral and the organic soil components. It therefore varies for different soils, e.g. within the group of mineral soils, and ranges from 2.4–2.9 g cm⁻³. Hence, awareness of this variability is important for properties estimated by a calculation involving particle density. Because ρ_S depends on both the soil's solid mineral particles and soil organic matter composition, we derived a function based on the mixture ratio of these two soil components. This approach represents a further development of earlier investigations dealing with the influence of organic carbon (C_org) on ρ_S. To parameterise this function, two data sets were used: (1) data from soils with C_org contents between 0% and 54.88% and corresponding values of ρ_S between 1.49 and 2.72 g cm⁻³; and (2) data from soils of 17 German long-term experiments contrasting in soil texture and in soil mineral inventory. Data set 1 was used to quantify the influence of soil organic matter on ρ_S, and data set 2 was used to calculate the influence of mineral matrix on ρ_S. The soil organic matter has two major influences on ρ_S: (1) via a mass effect (expressed as a mixture ratio between organic and mineral soil components); and (2) via a quality effect (expressed as calculated changes in particle density of organic soil components). Here, we calculated that with increasing content of soil organic matter (0–100%), the particle density of organic soil components rose from about 1.10 to 1.50 g cm⁻³, and present possible reasons for this phenomenon. Additionally, we demonstrate that the mineral matrix of the soil affects ρ_S especially via variations in the mineral inventory, but conclude that differences in particle size distribution of soils were to a lesser extent suitable for describing the influence of the mineral matrix on ρ_S. Overall, using our approach should generate more realistic values of ρ_S, and consequently of all calculated parameters which are sensitive to ρ_S.     

Phospholipid fatty acid profiles and xylanase activity in particle size fractions of forest soil and casts of Lumbricus terrestris L. (Oligochaeta,Lumbricidae)

Earthworms are key agents in organic matter decomposition, as they remove surface plant litter material and mix it with mineral soil. Plant litter material is comminuted in the gizzard of anecic earthworms and this is enhanced if sand particles are available. We hypothesize that this comminution of soil and litter will result in changes in the distribution of soil organic matter and soil microorganisms in the different particle-size fractions. We investigated soil organic matter content, xylanase- and microbial activity and community structure in bulk soil and particle size fractions of Lumbricus terrestris L. casts and in soil with and without the addition of beech litter.Earthworm gut passage did not affect the particle-size distribution but the content of soil organic matter was decreased in the fine sand fraction in treatments without litter (−6.80%) and increased in treatments with litter (+33.23%). The soil organic matter content of the clay fraction tended to be higher in earthworm casts. Xylanase activity was at a maximum in the fine sand fraction, lower in the coarse sand fraction and at a similar minimum in the silt- and clay-sized fraction. In the coarse sand fraction of the cast and litter treatments xylanase activity was increased by 39.1% and 124.8%, respectively. In the silt-sized fraction of casts the addition of litter increased xylanase activity (+58.6%) whereas, in casts without litter it was decreased (−36.25%). In the particle size fractions of casts, the content of bacterial PLFAs was decreased in the fine sand fraction and tended to be decreased in the clay fraction compared to the respective fractions in soil. In the silt fraction the fungal-to-bacterial PLFA ratio was higher in casts than in soil.We conclude that earthworms stabilize soil organic matter in cast aggregates predominantly by increasing the soil organic matter content in the clay fraction where it becomes protected against microbial attack. Organic matter in the coarse and fine sand fractions is decomposed primarily by fungi; xylanase is very active in these sand fractions and incorporation of litter into these fractions by the earthworms increased fungal biomass. Comminution of litter during passage through the earthworm gut increased the biomass and activity of fungi also in the silt fraction. The use of PLFA profiles in combination with other quantitative microbial methods improves the understanding of stabilizing and mobilizing processes in earthworm casts.