Changes in potassium availability and other soil properties due to soil ingestion by earthworms

An incubation experiment was conducted to study the changes that occur in potassium availability and other soil properties with ingestion of soil by earthworms. Two soils were used. Raumai soil with high non-exchangeable K and Milson soil with low non-exchangeable K were incubated with two species of earthworm, Aporrectodea caliginosa and Lumbricus rubellus, for 8 weeks. The casts and soil samples were analysed for exchangeable K, Ca, Mg, Na, and H, pH, organic C, and texture. The results indicated that in Raumai soil, the exchangeable K levels of the casts of both earthworm species were significantly higher than for the control soil, the effect being more marked for L. rubellus than for A. caliginosa. In Milson soil, the exchangeable K levels were significantly lower in the casts of both types of earthworm than in the control soil. The nitric acid-extractable K of the soil and casts was not markedly different for either soil type, but available non-exchangeable K values were significantly higher for the casts of L. rubellus from Milson soil than for the noningested Milson soil. In Raumai soil, the exchangeable Ca was higher in the casts of L. rubellus, exchangeable Mg and H were reduced, and exchangeable Na did not change markedly in the cast compared to the control soil. For Milson soil, the casts contained lower exchangeable Ca and H but higher Na and Mg than the control. The casts of both species of earthworm had significantly higher pH values for both soil types. There was no marked difference in the organic C content of the control soil and cast samples for Milson but a reduction in the casts of A. caliginosa for the Raumai soil. Finer fractions increased in the casts of both earthworm species in both soil types.     

The effect of earthworms (Lumbricidae) on nitrogenase activity in soil

Summary Nitrogenase activity associated with earthworms, their faeces and activity in soil was measured by the acetylene reduction technique. A clear increase in nitrogenase activity was found in field-deposited casts of Aporrectodea caliginosa in comparison with surrounding soil, although potential nitrogenase activity was significantly higher in soil than in casts. Nitrogenase activity associated directly with earthworms (Lumbricus rubellus ) was detected, indicating the presence of active N₂-fixing bacteria on the body surface and/or in the gut. Laboratory experiments showed that nitrogenase activity in the casts of L. rubellus was higher than in unmodified soil, and that nitrogenase activity in soil was significantly increased by the burrowing and feeding activity of these worms. This paper discusses the possible causes of these earthworm effects on soil nitrogenase activity and some methodological problems of determining the nitrogenase activity.     

Influence of soil ingestion by earthworms on the availability of potassium in soil: An incubation experiment

Summary An incubation experiment was conducted to study the changes that occur in the K status of soil due to earthworm activity. Samples of Tokomaru silt loam soil were inoculated with the common pasture earthworm species Aporrectodea caliginosa and incubated for 21 days. Aliquots of moist soil were analyzed for exchangeable K by leaching with neutral molar ammonium acetate at 1:50 soil solution ratio. Extraction with boiling 1 M nitric acid at 1:100 soil solution ratio for 20 min was used to determine available non-exchangeable K. The results indicated that the exchangeable K content increased significantly due to earthworm activity but nitric acid-extractable K did not change significantly. It is inferred that earthworms increase the availability of K by shifting the equilibrium among the forms of K from relatively un-available forms to more available forms in the soil chosen for the study.     

Population and behavioural level responses of arable soil earthworms to boardmill sludge application

The response of earthworms to soil application of boardmill waste sludge was quantified in field and laboratory experiments. The influence of one application of 6×10⁴ tonnes ha⁻¹ of unamended sludge was tested against no application on silty-clay arable soil. After 2 years, results in stubble-cultivated soil showed a 1.7 times lower density of earthworms where sludge was added, whereas there was no difference in average earthworm fresh weight. In direct-drilled soil, there was an indication of lower average numbers of Lumbricus terrestris middens when sludge was applied. Negative impacts in the field may have been due to indirect negative effects of the sludge application. In the laboratory, habitat choice tests were undertaken with two common species from the field site using sludge-based mulch and fertiliser products. Aporrectodea caliginosa showed no discernible preference between soil and soil mixed with unamended sludge (mulch), but favoured soil over soil mixed with sludge and chicken manure (fertiliser). Tests with L. terrestris showed a similar pattern. The preference of L. terrestris for the two products was compared with that for chopped barley straw by direct observation of foraging behaviour. Differences in foraging time of L. terrestris for different feeds were not significant, but mass of straw collected was significantly greater compared with either type of sludge treatment.     

Growth rates,establishment, and effects on herbage yield of introduced earthworms in grassland on reclaimed cutover peat

Summary Lumbricus terrestris L. juveniles confined in nylon mesh bags grew at mean rates of 6–12 mg ind^–1 day^–1 in reclaimed peat grassland soil, while the growth rates of Aporrectodea caliginosa (Sav.) juveniles were 1.5–2.1 mg ind^–1 day^–1. Earthworm population densities exceeding 700 m² had become established within 1 year adjacent to sods transplanted from an old pasture, while microplots enclosed in nylon mesh cages had mean population densities of 318–408 earthworms m^–2 and biomass of 89–111 g m^–2 3–4 1/2 years after inoculation. Herbage yields were 25% greater in the 2nd year and 49% greater in the 3rd year in earthworm-inoculated microplots which received an annual application of cattle slurry compared with similarly fertilized, non-inoculated cages.Dedicated to the late Prof. Dr. M.S. Ghilarov     

Ability of the earthworms Aporrectodea rosea and Aporrectodea trapezoides to increase plant growth and the foliar concentration of elements in wheat (Triticum aestivum cv. spear) in a sandy loam soil

In a greenhouse study, the ability of the earthworms Aporrectodea rosea and A. trapezoides to influence the foliar concentration of elements and the growth of wheat plants was assessed 27 days after sowing in a sandy loam soil. The presence of A. rosea and A. trapezoides (at densities equivalent to 314 and 471 m^-2, respectively) caused a significant increase in the shoot dry weight of wheat. The presence of A. rosea and A. trapezoides (at densities equivalent to 314 and 157 m^-2, respectively) was also associated with a significant increase in the root dry weight of wheat. The presence of A. rosea caused a significant increase in the foliar concentration of Ca, Cu, K, Mn, N, Na, and P, but did not influence the foliar concentration of Al, B, Fe, Mo, Mg, S, and Zn. The presence of A. trapezoides was associated with a significant increase in the foliar concentration of Al, Ca, Fe, K, Mn, N, and Na, but did not influence the foliar concentration of B, Cu, Mo, Mg, P, S, and Zn. These results demonstrate the potential of A. rosea and A. trapezoides to increase the growth of wheat in a sandy loam soil and suggest that the mechanism by which they increased plant growth was, in part, through increasing the availability and uptake of nutrients from this soil.     

Density, biomass and vertical distribution ofAporrectodea caliginosa (Savigny 1826) (Oligochaeta, Lumbricidae) in Benghazi, Libya

Studies were conducted during the period May 1993 to April 1994 on the density, biomass and vertical distribution ofAporrectodea caliginosa, a common species of earthworm found in the Benena farms in Benghazi, Libya. The soil of the farm was a clay loam, pH 7.3, and the organic matter averaged 4.2%. Maximum density and biomass ofA. caliginosa were recorded from October to December 1993 and again during March and April 1994, and minimum values were recorded from May to July 1993. A juvenile dominant population was observed during the summer. This was followed by an increase in subadults and adults during the autumn, winter and spring seasons. A positive correlation of the moisture and an inverse correlation of soil temperature on the density and biomass of these animals were discernible. A. caliginosa mainly confined their activities to the surface organic zone (0–10 cm depth) of the soil for most of the months studied.     

Effects of plant species on microbial biomass phosphorus and phosphatase activity in a range of grassland soils

Soil P transformations are primarily mediated by plant root and soil microbial activity. A short-term (40 weeks) glasshouse experiment with 15 grassland soils collected from around New Zealand was conducted to examine the impacts of ryegrass (Lolium perenne) and radiata pine (Pinus radiata) on soil microbial properties and microbiological processes involved in P dynamics. Results showed that the effect of plant species on soil microbial parameters varied greatly with soil type. Concentrations of microbial biomass C and soil respiration were significantly greater in six out of 15 soils under radiata pine compared with ryegrass, while there were no significant effects of plant species on these parameters in the remaining soils. However, microbial biomass P (MBP) was significantly lower in six soils under radiata pine, while there were no significant effects of plant species on MBP in the remaining soils. The latter indicated that P was released from the microbial biomass in response to greater P demand by radiata pine. Levels of water soluble organic C were significantly greater in most soils under radiata pine, compared with ryegrass, which suggested that greater root exudation might have occurred under radiata pine. Activities of acid and alkaline phosphatase and phosphodiesterase were generally lower in most soils under radiata pine, compared with ryegrass. The findings of this study indicate that root exudation plays an important role in increased soil microbial activities, solubility of organic P and mineralization of organic P in soils under radiata pine.     

Significance of earthworms in stimulating soil microbial activity

 The stimulatory effect of earthworms (Lumbricus terrestris L.) on soil microbial activity was studied under microcosm-controlled conditions. The hypothesis was tested that microbial stimulation observed in the presence of a soil invertebrate would be due to the utilization of additional nutritive substances (secretion and excretion products) that it provides. Changes in microbial activity were monitored by measuring simultaneously CO₂ release and protozoan population density. The increase in CO₂ released in the presence of earthworms was found to result from both earthworm respiration and enhanced microbial respiration. The stimulation of microbial activity was confirmed by a significant increase in protozoan population density, which was 3–19 times greater in the presence of earthworms. The respiratory rate of L. terrestris was estimated to be 53 μl O₂ g^–1 h^–1. Earthworm respiration significantly correlated with individual earthworm weight, but there was no correlation between the increase in microbial respiration and earthworm weight. This finding does not support the hypothesis given above that enhanced microbial respiration is due to utilization of earthworm excreta. A new hypothesis that relationships between microbial activity and earthworms are not based on trophic links alone but also on catalytic mechanisms is proposed and discussed. Received: 26 August 1997     

Partitioning of ryegrass residue sulphur between the soil microbial biomass,other soil sulphur pools and ryegrass (Lolium perenne L.)

Summary Ryegrass shoot residues, labelled with ³⁵S, were added to an S-deficient soil. The transfer of S to the microbial biomass, to the soil S pool extractable by NaHCO₃ and to growing ryegrass when present was followed over 34 weeks. After 2 weeks 16% and 15% of the S residue was found in the biomass and in the extractable S pool, respectively. Where plants were grown, they became S-deficient (shoot S <0.2%) simultaneously with the biomass showing a marked increase in C:S ratio. This eventually reached 262 from an initial value of 59. Concurrently, the extractable S pool, which included some labile organic S, decreased to <0.2 g g^–1 soil. After 34 weeks 27% of the S residue was found in the growing plant, 7% in the biomass and 2% in the extractable S pool. Some mineralization of unlabelled soil organic S was observed during the period of greatest plant growth (8–14 weeks), but not in the absence of plants. A second phase of mineralization occurred between weeks 22 and 34, concurrent with a rise in mean temperature, which was unaffected by the presence of plants or by the size of the microbial biomass. This may have been due to biochemical mineralization of ester sulphate. The amount of unlabelled soil S involved in active cycling was estimated to be 11%–13% of the total soil S.     

Mucus excretion and carbon turnover of endogeic earthworms

Summary Mucus excretion of endogeic earthworms, by the body surface to burrow walls and by the intestine to casts, was investigated using uniformly ¹⁴C-labelled adult or subadult specimens of Octolasion lacteum (Örley) in laboratory incubations in soil from a beechwood on limestone. The daily loss of C due to mucus excretion from the body surface and in casts was calculated as 0.2 and 0.5% of total animal C, respectively. The C loss due to mucus excretion by subadult or adult individuals of O. lacteum is assumed to account for 63% of total C losses (including mucus excretion and respiration) of the earthworms. In a second experiment we studied the incorporation of ¹⁴C from labelled soil, again from a beechwood on limestone, into the tissue of the endogeic earthworm species Aporrectodea caliginosa (Savigny). The results of this experiment indicate the existence of two C pools, one more labile and one more stable, in earthworms. It is assumed that the C investment for respiration and mucus excretion is derived from the labile C pool of endogeic earthworms.     

Weight response to the soil water potential of the earthworm Aporrectodea longa

Summary To assess the ability of endogeic earthworms to tolerate variations in soil water potential, groups of these worms were subjected to different, constant levels of soil water suction () over a period of 17 days. At water suctions varying from 0.3 kPa (pF 0.5) to 1990 kPa (pF 4.3), the earthworms showed no physiological ability to a maintain constant internal water content as assessed by the fresh weight. The relationship between weight loss and the increase in water, suction was modelled and the following critical thresholds were identified: <60 kPa did not affect earthworm weight; >620 kPa led to diapause; between these two values, earthworm weight was closely governed by the variation in . At the intermediate suction of 167 kPa, the exchange of water between the earthworms and the soil was at a maximum, and this value is therefore proposed as the level at which the diapause is induced. These modelled values were the diapause is induced. These modelled values were compared with those obtained under field conditions.     

Growth rates of Aporrectodea caliginosa (Oligochaetae: Lumbricidae) as influenced by soil temperature and moisture in disturbed and undisturbed soil columns

Earthworm growth is affected by fluctuations in soil temperature and moisture and hence, may be used as an indicator of earthworm activity under field conditions. There is no standard methodology for measuring earthworm growth and results obtained in the laboratory with a variety of food sources, soil quantities and container shapes cannot easily be compared or used to estimate earthworm growth in the field. The objective of this experiment was to determine growth rates of the endogeic earthworm Aporrectodea caliginosa (Savigny) over a range of temperatures (5–20 °C) and soil water potentials (−5 to−54 kPa) in disturbed and undisturbed soil columns in the laboratory. We used PVC cores (6 cm diameter, 15 cm height) containing undisturbed and disturbed soil, and 1 l cylindrical pots (11 cm diameter, 14 cm height) with disturbed soil. All containers contained about 500 g of moist soil. The growth rates of juvenile A. caliginosa were determined after 14–28 days. The instantaneous growth rate (IGR) was affected significantly by soil moisture, temperature, and the temperature×moisture interaction, ranging from −0.092 to 0.037 d⁻¹. Optimum growth conditions for A. caliginosa were at 20 °C and −5 kPa water potential, and they lost weight when the soil water potential was −54 kPa for all temperatures and also when the temperature was 5 °C for all water potentials. Growth rates were significantly greater in pots than in cores, but the growth rates of earthworms in cores with undisturbed or disturbed soil did not differ significantly. The feeding and burrowing habits of earthworms should be considered when choosing the container for growth experiments in order to improve our ability to extrapolate earthworm growth rates from the laboratory to the field.     

Microbial activity and nutrient dynamics in earthworm casts (Lumbricidae)

Summary Microbial respiration, microbial biomass and nutrient requirements of the microflora (C, N, P) were studied in the food substrate (soil taken from the upper 3 cm of the mineral soil of a beech wood on limestone), the burrow walls and the casts of the earthworm Aporrectodea caliginosa (Savigny). The passage of the soil through the gut caused an increase in soil microbial respiration of about 90% over a 4-week period. Microbial biomass was increased only in freshly deposited casts and decreased in aging faeces to a level about 10% lower than in soil. Microbial respiration of the burrow walls was only increased over a shorter period (about 2 weeks). The microflora of the soil and the burrow walls was limited by P, whereas in earthworm casts, microbial growth was limited by the amount of available C. In aging faeces the P requirement of the microflora increased and approached that of the soil. Immobilization of phosphate in earthworm casts is probably caused by mainly abiotic processes. C mineralization by soil microflora fertilized with glucose and P was limited by N, except in freshly deposited casts. Ammonium, not nitrate, was responsible for this process. N dynamics in earthworm casts are discussed.     

Effects of endogeic earthworms on soil processes and plant growth in coniferous forest soil

Summary The effects of the endogeic earthworm, Aporrectodea caliginosa tuberculata (Eisen) on decomposition processes in moist coniferous forest soil were studied in the laboratory. The pH preference of this species and its effects on microbial activity, N and P mineralization, and the growth of birch seedlings were determined in separate pot experiments. Homogenized humus from a spruce stand was shown to be too acid for A. c. tuberculata. After liming, the earthworms thrived in the humus and their biomass increased (at pH above 4.8). In later experiments in which the humus was limed, the earthworms positively influenced the biological activity in humus and also increased the rate of N mineralization. A. c. tuberculata increased the growth of birch seedlings, with increases observed in stems, leaves, and roots. Neither NH ₄ ⁺ -N fertilizer nor mechanical mixing with artificial worms affected seedling growth. No plant-growth-affecting compounds (e.g., hormone-like compounds) due to the earthworms were present in the humus. The shoot: root ratio in the birch seedlings was not affected by either the earthworms or the fertilizer. The experiments revealed the impact of earthworm activity on soil processes and plant growth.     

Burrowing ability of the earthworm Aporrectodea longa limited by soil compaction and water potential

Summary Adult earthworms (Aporrectodea longa) were maintained for 199 days in soil columns (h=30 cm; ø=10 cm) where the water potential ranged from -7 to-65 kPa and compaction from 50 to 350 kPa. The weight of casts on the soil surface was measured at the end of two periods of activity (spring and autumn). Cast production increased with bulk density, but the activity of earthworms was limited both by the mechanical strength of the soil and by decreasing water potential. The results obtained in the laboratory conditions of this study were consistent with field observations on casting and burrowing activities. The effects of water potential and soil compaction on these activities were estimated.     

Cropping history and peat amendment-induced changes in strawberry field earthworm abundance and microbial biomass

Common agricultural practices, e.g. soil tillage and organic amendment, may affect field earthworm communities considerably. However, there is little data to show how long the changes persist after a certain action. The effect of peat, commonly used in Finland to improve the horticultural soil structure, on key soil organisms is also largely unknown. Earthworm abundance and microbial biomass were studied in a strawberry field experiment (soil type silty clay) with a history of different crops (strawberry, timothy, caraway, rye, turnip rape, fiddleneck, onion and buckwheat) and peat treatments. Sampling was carried out after three years of perennial cropping of strawberry. Half of the area was peat-amended twice three years apart. The earthworm community consisted mainly of Aporrectodea caliginosa and Lumbricus terrestris. Soil peat amendment almost doubled the number of endogeic A. caliginosa, but had no effect on the anecic L. terrestris. The effect of cropping history on earthworms diminished after three years of strawberry cropping. Only the positive effect of caraway on juvenile Lumbricus spp. was detectable three years after its cropping had been finished. However, some crops had secondary effects on the earthworm distribution without significant influence on their numbers while they were grown, e.g. high numbers of A. caliginosa were recorded from soil with a history of timothy ley. The effect of strawberry cropping was contradictory: six years of continuous strawberry cropping decreased the number of the anecic L. terrestris, but during the last three years on strawberry, the proportion of L. terrestris increased from 6% to 40% in the experimental area with a concomitant great drop in the number of A. caliginosa. The role of different agricultural practices (no tillage, mulching, inter-row grass cover and pesticides) is discussed. The crop-induced changes persisted in the microbial biomass for three years (onion cropping reduced microbial biomass C), but soil amendment had no effect on microbes. The abundance of A. caliginosa was associated with soil organic C, but not with soil microbial biomass.