Mechanisms of stabilization of earthworm casts and artificial casts

Summary Fresh casts were collected from the earthworm species Aporrectodea caliginosa, and artificial casts were also made. The casts were subjected to ageing, drying-rewetting, and sterilization by hexanol vapour. Clay dispersion was determined, as a measure of the lack of stability of the casts. Two soils were used, the topsoil of a recently reclaimed polder soil in the Netherlands and the topsoil from a South Australian duplex soil. For both soils the fresh worm casts had higher dispersible clay than the artificial casts. During ageing, both types of casts became more stable. There are strong indications that this was mainly due to changes on the surface of the casts. Fungi developed on the surface of 6-day-old worm casts made of Australian soil. This gave a higher stability to the casts compared to artificial casts of the same age without fungal growth. With both types of casts, hexanol inhibited fungal growth on the surface of the casts, reducing the stabilizing effect of ageing. The fungus did not develop on Dutch soil casts until after 42 days, and the development of a higher stability with age was also less marked than in the Australian soil. When the casts were subjected to a drying and rewetting cycle before analysis, they became much more stable than the casts that were analyzed wet. The drying-rewetting cycle removed most of the differences between the worm casts and the artificial casts, and also removed any effects of ageing.Dedicated to the late Prof. Dr. W. Kühnelt     

Protection of soil carbon by microaggregates within earthworm casts

Earthworms are known to play a role in aggregate formation and soil organic matter (SOM) protection. However, it is still unclear at what scale and how quickly earthworms manage to protect SOM. We investigated the effects of Aporrectodea caliginosa on aggregation and aggregate-associated C pools using ¹³C-labeled sorghum (Sorghum bicolor (L.) Moench) leaf residue. Two incubations were set up. The first incubation consisted of soil samples crushed <250 μm to break up all macroaggregates with three treatments: (i) control soil; (ii) soil+¹³C-labeled residue and (iii) soil+¹³C-labeled residue+earthworms. Earthworms were added after 8 d and 12 d (days) later, aggregate size distribution was measured together with total C and ¹³C in each aggregate fraction. A second incubation was made to assay protected versus unprotected total C and ¹³C from 21-d laboratory incubations of intact and crushed large (>2000 μm) and small (250-2000 μm) macroaggregates and microaggregates (53-250 μm). Eight different pools of aggregate-associated C were quantified: (1) and (2) unprotected C pools in large and small macroaggregates, (3) unprotected C pools in microaggregates, (4) and (5) protected C pools in large and small macroaggregates, (6) protected C pool in microaggregates, and (7) and (8) protected C pools in microaggregates within large and small macroaggregates. In the presence of earthworms, a higher proportion of large macroaggregates was newly formed and these aggregates contained more C and ¹³C compared to bulk soil. There were no significant differences between the samples with or without earthworms in the C pool-sizes protected by macroaggregates, microaggregates or microaggregates within small macroaggregates. However, in the presence of earthworms, the C protected by microaggregates within large macroaggregates was a significant pool and 22% of this C pool was newly added C. In conclusion, these results clearly indicate the direct involvement of earthworms in providing protection of soil C in microaggregates within large macroaggregates leading to a possible long-term stabilization of soil C.     

Nutrient content of earthworm casts influenced by different mulch types

The aim of this study was to investigate the effects of different mulch types on the quality and quantity of casts of different lumbricid earthworm species. Nutrient contents (organic carbon C_org, total nitrogen N_t and inorganic phosphate Pi), and acid phosphatase activity (SPA) of earthworm casts and soil aggregates were measured and then compared. This short time experiment was conducted in the laboratory, simulating field conditions of mulch management in temperate agroecosystems. In microcosms, the two common field species Lumbricus terrestris and Octolasion cyaneum were inoculated. Barley, lupin, maize, or sugarbeet were used as mulch according to amounts usually applied in the field. Nutrient contents and phosphatase activity in worm casts were generally enhanced compared to the soil. Nutrient values were higher in the casts of the detritivorous L. terrestris than those of the geophageous O. cyaneum. Conversely, the phosphatase activity was increased in the casts of O. cyaneum compared to L. terrestris in case of the barley and sugar-beet treatments. Cast production was related to the earthworms' change of biomass (ΔB) as well as, in the case of L. terrestris, to nitrogen content of the mulch. Due to their chemical compounds, the mulch types influence both the food selection of the worms and the acceptance by microorganisms.     

An evaluation of 2D-image analysis techniques for measuring soil microporosity

To compare soil porosity measurements, manual, semiautomatic and automatic 2D‐image analyses were performed on three sets of images of the same fields. The first and second image sets were obtained, by a fluorescence microscope, on the polished surfaces of soil blocks impregnated with a fluorescent resin and on the thin sections made from them, respectively. A scanning electron microscope in backscattered mode was used to acquire the third set of images on the thin sections. In the manual image analysis procedure, image segmentation was based on the best visual impression and carried out individually for each image with the UTHSCSA Image Tool software. For the semiautomatic method, the software μCTanalySIS was used; the images were segmented by double hysteresis, after interactive selection of the thresholding values. Automatic thresholding, based on an analysis of the image intensity histogram, was performed using the image analysis software Image‐Pro^® Plus. Average microporosity measurements were compared between image types and image analysis methods using marginal regression models for continuous outcomes. The mean area covered by pores was significantly different depending on the type of image and the method of image analysis. Results after the automatic procedure were significantly larger than after manual or semiautomatic thresholding. On the other hand, average porosities were larger for backscattered electron images than for fluorescent images, regardless of the image analysis procedure used. These results stress the need for standardization of image analysis protocols and warn of the dangers of comparing soil porosity measurements performed on different types of images.     

The interactions between soil type and earthworm species determine the properties of earthworm casts

Earthworms are recognized to increase soil porosity, reorganize soil structure, and stimulate soil microflora and nutrient mineralization. The properties of earthworm casts should depend both on earthworm species or ecological group and on soil properties. Interactions between earthworm species and soil types have been suggested, but only poorly demonstrated. In order to better understand those interactions, two hypotheses led our study: (1) Soil type has a greater influence on cast properties than earthworm; (2) Earthworms from different species influence cast properties differently; (3) The intensity and direction of the impact of each earthworm species on cast properties vary with soil properties. Fifteen physical and chemical variables (N–NH₄⁺, N–NO₃⁻, total organic C and N, C/N ratio, CaCO₃, pH, P, K⁺, Mg²⁺, Mn²⁺, Na⁺, CEC, moisture, wettability) were measured in casts of three earthworm species (Lumbricus terrestris, Allolobophora chlorotica and Aporrectodea rosea) produced in three temperate soils. Univariate and multivariate analyses showed that earthworm species and soil types significantly impacted cast properties. pH, N_t, K and Mg contents were interactively altered by both factors. Multivariate analysis showed that a difference of soil type had a major impact on casts properties (62%) compared to the impact of a difference of earthworm species (10%). Cast properties were most impacted by L. terrestris, then by A. chlorotica and last by A. rosea. The response ratio (ratio of the properties of the casts to the properties of the bulk soil) was used to quantify the effect of earthworm species compared to the control soil. It showed a higher response of variables in casts in nutrient-rich soils, especially in casts of L. terrestris. The interactions between earthworm species and soil types on cast properties were discussed with regards to earthworm ecology, properties of the soil, and earthworm modifications of cast microflora.     

Characteristics of earthworm casts affecting herbicide sorption and movement

Various physical and chemical characteristics of earthworm casts collected from a laboratory incubation and a field experiment were examined in relation to their effect on the sorption and the movement of three ¹⁴C-labelled ionic herbicides: atrazine, 2,4-dichlorophenoxyacetic acid, and metsulforon methyl. The earthworm casts contained higher levels of fine fractions and total and soluble C. This is attributed to the grinding action of the earthworm gut and selective feeding on zones with higher organic matter and fine size fractions. The earthworm casts had a higher pH than the source soil, resulting in a higher number of surface negative charges. The earthworm casts sorbed higher amounts of herbicides than the source soil, mainly due to the increases in the amount of organic C and fine size fractions. The incrased sorption of herbicides by the casts resulted in decreased leaching.     

Effect of earthworm casts on protein synthesis inAgaricus bisporus

Summary When earthworm casts were used as a casing, an early fructification, a greater stimulation in fruitbody formation, an increase in the carpophore protein content and in L-¹⁴C-leucine incorporation were observed inAgaricus bisporus. The protein-synthesizing capacity of the carpophore and its protein content were tested on samples collected 28 and 35 days after the casing. An increased L-¹⁴C-leucine incorporation of about 34% and an increased protein content of about 25% were recorded.     

The hormone-like effect of earthworm casts on plant growth

Summary The fertilizing effect of earthworm casts depends on microbial metabolites, mainly growth regulators. The hormone-like effect of earthworm casts is discussed with reference to the literature and ad hoc experiments. When used in plant propagation, earthworm casts promote root initiation and root biomass and increase root percentage. When applied as a casing layer, earthworm casts stimulate carpophore formation in Agaricus bisporus, and N assimilation. When used in horticulture, earthworm casts have a hormone-like effect, influencing the development and precociousness of plants or inhibiting them. These effects are dependent on dose, application time and plant species. In addition, results recorded on dwarfing, stem elongation and precociousness of flowering suggest that the biological effect of earthworm casts is linked to microbial metabolites that influence plant metabolism, growth and development.     

Rainfall effects on erosion of earthworm casts and phosphorus transfers by water runoff

 We investigated whether, under a temperate climate and in a maize crop, earthworm casts could contribute to soil erosion and further favour the exportation of phosphorus by runoff waters. Recording of casts was made in compacted (wheel-tracks) and non-compacted inter-rows, for a 2-month period in spring. To assess the rainfall impact on cast evolution, half of the observation sites were protected against rain splash by a nylon mesh placed above the soil surface. The water runoff was collected and analysed for sediment contents and phosphorus concentration. The mean annual production of surface casts was calculated to be 34 kg (dry weight) year^–1 kg^–1 earthworm (fresh weight). Synchronization between cast erosion and rainfall events was shown under natural conditions (unprotected sites). The erosion rate was 4 times greater over rainy periods than dry ones, reaching 80% of cast numbers. It appeared that not the runoff effect but the splash effect, due to the kinetics of the drops, disrupted casts. Newly formed casts disappeared first, with the erosion rate decreasing twofold for casts more than 10 days old. Cast erosion and runoff, as well as worm casting activity, were greater under compacted sites than under non-compacted sites, indicating an influence of earthworms on soil erosion from compacted soils. The total phosphorus content was similar in casts and uningested soil (0.80 mg phosphorus g^–1). Potential phosphorus losses from cast erosion was calculated to reach 25–49 mg phosphorus m^–2 per rainfall event depending on soil compaction. The amounts of particulate phosphorus recovered in water runoff after each rainfall event varied from 1 mg to 11 mg phosphorus. These results are compared and discussed. Received: 20 October 1998     

Rainfall impact effects on ageing casts of a tropical anecic earthworm

We investigated the erodibility of surface casts produced by an anecic earthworm of the Colombian savannahs by means of indoor rain simulations. The kinetic energy applied to samples, 21.62 J minute⁻¹ m⁻², was estimated to be equivalent to 41% of the energy of the more intense period of a local storm. The erodibility of casts was assessed at different stages of their ageing along with the effects of repeated wetting‐drying cycles. Bare soil cores and soil cores of the same size with a cast on their surface (soil + cast) were used as controls. Saturated hydraulic conductivity (HC) was measured to test whether casts enhance soil water permeability. Fresh, almost liquid, individual casts were completely dispersed by a 2‐hour rainfall simulation at a dispersion rate of 0.9 ± 0.5% of sample minute⁻¹. After 5 hours drying at 32°C and 79% relative humidity, casts were as moist as fresh casts (non‐significant Mann–Whitney U‐test) but had a solid appearance and were no longer dispersed by raindrop impact. Under simulated rainfall, dry casts were very slowly fragmented into large aggregates (> 5 mm). The HC of casts was increased by repeated cycles of wetting and drying but unaffected by length of air‐drying and natural ageing, and averaged 7 (3, n = 6), 20 (3, n = 7) and 32 (1, n = 45) cm hour⁻¹ for bare soil, soil + cast and all casts, respectively (standard error, sample size). Except for bare soil, these values were greater than the greatest rainfall intensities recorded locally. The consequences for soil erosion, nutrient losses and water infiltration are discussed.     

Potential role of earthworm casts for the phosphorus enrichment of run-off waters

Surface casts of earthworms (dominantly Allolobophora caliginosa) contained a higher proportion of finer particles than the underlying surface soil. The proportion of the finest fraction (< 4 μm), recovered following water dispersion, increased from 6% in the surface soil to 20% in casts. Casts contained appreciably more inorganic and organic phosphorus (P) than underlying soil materials, particularly in the finer fractions (20-4 and < 4 μm). The release of inorganic P to solution from casts was approximately four times greater than that from surface soil. In contrast, the removal of inorganic P from solution by casts was proportionately lower than that by surface soil. Studies of the rate of release of inorganic P to solution indicated the presence in the casts of a pool of loosely-bound inorganic P which was readily released to solution. This was confirmed by isotopic exchangeability studies, the amount of exchangeable P being three times greater in casts than in surface soil. Although the release of total P to solution from fresh casts did not change with time, the release of inorganic P increased as a function of time of incubation. Data for phosphatase activity in casts indicated that 0.1 m NaCl-extractable organic P was converted to inorganic P and that this was quantitatively released to solution. The results obtained are discussed in terms of the potential contribution of surface casts to the particulate and dissolved P loadings of run-off waters.     

Degradation dynamics of surface earthworm casts in grasslands of the eastern plains of Colombia

 Earthworms are generally considered to fit the definition of ecosystem engineers. The casts they produce are recognised to have a great importance in the regulation of soil processes. Lifetimes and degradation rates of these structures remain poorly known. In this study, the dynamics of disappearance and the changes in the physical properties of the surface casts of the anecic earthworm Martiodrilus carimaguensis were assessed in a native savanna and an intensive pasture. In both systems, casts were composed of superposed layers deposited by earthworms over a period of at least a few days. The half-life of casts ranged between 2 months and 11 months in the pastures (trampled and protected, respectively), and 5 months in the savanna. Some dry casts remained at the soil surface for more than 1 year after having been excreted. The disappearance of casts was mostly attributed to rain-drop impact and the effect of animal trampling. The bulk density of fresh casts was higher (+17%, P<0.05) or equivalent (–1%, P>0.05) to that of the surrounding soil, in the savanna and the pasture, respectively. Fresh cast aggregates were of larger size than bulk soil aggregates (about +70% in both systems, P<0.05). Bulk density and the size of cast aggregates decreased with cast ageing (from –29% to –24% for bulk density, and from –68% to –80% for size, in the pasture and the savanna, respectively). Macroinvertebrates were observed digging into casts, and were assumed to be partly responsible for the physical degradation of casts with time. Received: 12 November 1999     

Destabilization of soil during the production of earthworm (Lumbricidae) and artificial casts

The mechanisms by which soil is destabilized in the digestive tract of endogéic earthworms were investigated with artificial casts, which were moulded with a syringe from slurries of a silty loam with or without gypsum and organic matter treatments, and compared to casts produced by Aporrectodea rosea (Lumbricidae). Both types of casts generally had the same levels of mechanical dispersion, observations of slaking, and particle size distribution when the casts were fresh, aged-moist for 30 days, or air-dried. Fresh casts were significantly more dispersive than the uningested soil despite the addition of gypsum or organic matter to the soil. However, the dispersion from aged-moist or air-dried casts was not substantially greater than that of uningested soil. Air-drying was more effective than moist-ageing in increasing the stability of casts and uningested soil. The concentration of soluble carbohydrate was greater in artificial casts produced from soil treated with sheep dung or xanthan gum, and in earthworm casts produced from soil treated with xanthan, than in the uningested soil of the same treatments. An increase in the concentration of soluble carbohydrate was related to an increase in dispersion. An attempt was made to simulate the addition of mucus to soil in the digestive tract of earthworms, by the addition of sucrose or xanthan gum to the slurry during the production of artificial casts. The addition of xanthan, but not sucrose, to the slurry increased mechanical dispersion relative to that of the uningested soil in the fresh treatment. Although the production of artificial casts destabilized soil to the same degree as earthworm casts, the artificial casts did not simulate all chemical, biochemical, and microbiological aspects of digestion. Received: 24 November 1995     

Influence of drying and ageing on the stabilization of earthworm (Lumbricidae) casts

The influence of drying and ageing on the stabilization of casts produced by the endoge′ic earthworm, Aporrectodea rosea, from a soil, which was hard-setting and low in organic matter, were investigated in the laboratory. Casts and uningested soil were aged-most for up to 32 days, dried for up to 21 days, or subjected to different wetting and drying cycles over 30 days. The dispersion index of aged-moist casts decreased from 0.40 to 0.25 over 32 days, while dispersion index of dried casts decreased from 0.40 to 0.01 over 21 days. The dispersion index of air-dried casts was not significantly increased by five cycles of wetting and drying. The dispersion index of dried casts was not significantly less than that of dried soil. In soils wetter than a matric potential of approximately –35kPa, stabilization of casts was probably due to a combination of cohesion of soil particles, age-hardening and growth of microorganisms. However, in soils drier than –35kPa, cementation was probably the major mechanism of stabilization. The addition of wheat straw to the soil prior to ingestion by earthworms increased dispersion from aged-moist casts, but did not influence dispersion from dried casts. The addition of wheat straw decreased the number of air-dried casts which slaked severely. The concentration of soluble carbohydrate decreased with dispersion index as casts and uningested soil were each dried. This suggested that soluble carbohydrate may have been denatured with or without being bonded to soil particles during drying. Received: 7 May 1996     

Effects of fallow system and cropping frequency upon quantity and composition of earthworm casts

The quantities of earthworm surface casts were monitored in traditional bush fallow regrowth (BFR), Pueraria phaseoloides live mulch (PLM), Leucaena leucocephala alley cropping (LAC) and undisturbed forest. The fallow systems were planted to maize/cassava intercrop either permanently or for one year after three years of fallow i.e. at 100 and 25% cropping frequencies. Earthworm casting activity was lower in fields cropped after clearing three year old fallow than in the respective permanently cropped fallow management system. The reduction in casting was related both to the degree of biomass removed through burning and to re-establishment of cover crops. Higher exchangeable cation concentration in the soil did not cause increased casting activity. Concentrations of organic C, total N and exchangeable Ca and Mg in casts were significantly higher than in the 0–15 cm topsoil. Exchangeable Ca and Mg in casts did not significantly differ between treatments, with the exception of significantly higher Mg in casts after three years of P. phaseoloides fallow. Topsoil (0–15 cm) exchangeable Ca and Mg were not correlated with cast exchangeable Ca and Mg, but concentrations of organic C and total N in casts were significantly correlated with organic C and total N in the topsoil. Coefficients of variability of organic C, total N and exchangeable Ca and Mg were about twice as high in the soil than in casts. Significant negative correlations between the cast to soil ratio (cast enrichment factor) of organic C, total N and exchangeable Ca and Mg and the respective concentrations in the soil show that earthworms are increasingly selective in organic carbon and nutrient uptake as these parameters of soil fertility are declining.     

Effect of agroforestry woody species on earthworm activity and physicochemical properties of worm casts

The effects of five agroforestry woody species (Dactyladenia barteri, Gliricidia sepium, Leucaena leucocephala, Senna siamea andTreculia africana) on the surface aasting activity ofHyperiodrilus africanus were studied in an Alfisol (Oxic Paleustalf) in southwestern Nigeria. Casting activity under the woody species decreased in the following order:Dactyladenia sp. (26.4 Mg ha^-1 year^-1)>Gliricidia sp. (24.4 Mg ha^-1 year^-1)>Treculia sp. (22.9 Mg ha^-1 year^-1)>Leucaena sp. (18.6 Mg ha^-1 year^-1)>Senna sp. (18.3 Mg ha^-1 year^-1). These differences in casting activity were partly explained by microclimatic effects. Irrespective of the woody species, the worm casts were higher in clay and silt contents, bulk density, water-stable aggregates, pH, organic C, exchangeable cations, effective cation exchange capacity, and extractable P levels than the corresponding surface soils. The woody species did affect the physicochemical properties and P sorption of the worm casts. The content of water-stable aggregates of worm casts decreased in the following order:Dactyladenia sp.>Treculia sp.>Senna sp.>Leucaena sp.>Gliricidia sp. Large differences in extractable P levels were observed.Senna sp. was associated with the highest extractable P level (11.5 mg kg^-1) andTreculia sp. the lowest (4.9 mg kg^-1). P sorption was highest on worm casts underDactyladenia sp. and lowest on those underTreculia sp. Without fertilizer application, there were no significant differences in the dry weight of maize grown in the different worm casts. With NPK applications, the dry weight of maize grown in worm casts associated withTreculia sp. was significantly lower than that of maize grown in the other worm casts, mainly due to the low extractable P level. Despite a high organic C and exchangeable K status, maize grown in the worm casts still responded significantly to N and K applications. The N uptake by maize grown in worm casts associated withTreculia sp. was lower than that in the other treatments.     

Effect of biosolid amendment on enzyme activities in earthworm (Lumbricus terrestris) casts

We studied the effect of amendment of sewage sludge biosolids on enzyme activity in soil and earthworm (Lumbricus terrestris) casts. Enzyme activities and contents of nutrients and organic matter of surrounding soil were compared with the corresponding properties of earthworm casts. This short time experiment was conducted at 20 ± 0.5 °C in the laboratory, simulating field conditions of biosolid treatments. In general, all of doses of biosolid treatments influenced the enzyme activity and contents of nutrients and organic matter of earthworm casts and surrounding soil. Enzyme activity such as urease (UA), alkaline phosphatase (APA), and arylsulfatase (ASA) and the contents of organic matter and nutrients N and P in earthworm casts and surrounding soil increased with increasing biosolid application. Without biosolid additions, enzyme activities in cast of L. terrestris exceeded those in the soil. In contrast, when biosolid was added, DHA in casts was lower than the soil. Activities of UA and APA were consistently higher in L. terrestris casts than in soil of all biosolid treatments. Biosolid amendments generally increased ASA at low doses, but at higher doses, ASA decreased. In general, organic matter and contents of N and P were higher in surface casts of L. terrestris and soils than in the control soil. Activities of UA, APA, the contents of organic carbon and nutrients N and P in soil and casts showed positive correlations. On the contrary, ASA and DHA were negatively correlated with the contents of organic matter and nutrients.     

Potential for higher rates of denitrification in earthworm casts than in the surrounding soil

Summary Earthworms (Lumbricus terrestris L.) were cultured in the laboratory and fed on lucerne (Medicago sativa L.). Denitrification rates in the surface casts and the surrounding soil were quantified using C₂H₂-inhibition of nitrous oxide reductase. The investigation also included determination of the N₂O-formation by nitrification as well as CO₂-formation as a measure of respiration. The denitrification rates of wet earthworm casts were found to be significantly higher than those occurring in wet samples from the soil. The low N₂O-formation observed seemed to be due to denitrification. Respiration was higher in casts, indicating higher oxygen demand which resulted in more anaerobic conditions. The energy supply was probably better in casts compared with the surrounding soil.