Impact of the earthworm Lumbricus terrestris on the degradation of Fusarium-infected and deoxynivalenol-contaminated wheat straw

When conservation tillage is practised in agriculture, plant residues remain on the soil surface for soil protection purposes. These residues should be widely decomposed within the following vegetation period as microbial plant pathogens surviving on plant litter may endanger the currently cultivated crop. Important soil-borne fungal pathogens that preferably infect small grain cereals belong to the genus Fusarium. These pathogens produce the mycotoxin deoxynivalenol (DON), a cytotoxic agent, in infected cereal organs. This toxin frequently occurs in cereal residues like straw. So far it is unclear if DON degradation is affected by members of the soil food web within decomposing processes in the soil system. For this purpose, a microcosm study was conducted under controlled laboratory conditions to investigate the degradation activity of the earthworm species Lumbricus terrestris when exposed to Fusarium-infected wheat straw being contaminated with DON.Highly Fusarium-infected and DON-contaminated straw seemed to be more attractive to L. terrestris because it was incorporated faster into the soil compared with straw infected and contaminated at low levels. This is supported by a greater body weight gain (exposure time 5 weeks) and smaller body weight loss (exposure time 11 weeks) of L. terrestris, respectively, when highly contaminated straw was offered for different time periods.Furthermore, L. terrestris takes part in the efficient degradation of both Fusarium biomass and DON occurring in straw in close interaction with soil microorganisms. Consequently, earthworm activity contributes to the elimination of potentially infectious plant material from the soil surface.     

Genetic structure among earthworms (Lumbricus terrestris L.) from different sampling sites in western Germany based on random amplified polymorphic DNA

Earthworms are being used as bio-indicators to assess terrestrial pollution. However, it is often not known whether their populations possess a uniform genetic structure, which would allow comparison of residues or biological properties of earthworms from different sampling locations. In order to investigate this point, random amplified polymorphic DNA (RAPD) variation was surveyed in earthworms (Lumbricus terrestris) from five different sampling sites in Germany. Forty oligonucleotide RAPD primers (10 base pairs in length) were screened, three of which produced high polymorphic band patterns. A total of 61 DNA fragments were detected in 90 individuals of L. terrestris from five sampling sites with 49 (80.3%) RAPD markers being polymorphic. The genetic similarities within (band sharing rates between 0.756 and 0.795) and among the L. terrestris populations (0.635) were similar even at widely separate locations. Inter-population variation in the RAPD pattern for all five earthworm populations accounted for 37.9% of the total variation, while intra-population variation for three adjacent Saarland populations accounted for only 18.0% of the total variation. Principal component analysis (PCA) and the genetic distances of the populations confirm these results. Twenty-four percent of the genetic distance is caused by geographical isolation as shown by a test for isolation by distance. These results show that L. terrestris fulfils the genetic qualifications for a bio-indicator particularly at closely located sampling sites. However, the results also suggest that earthworm studies of widely separated locations should include genetic characterisation of the earthworm samples.     

Quantitative estimates of burrow construction and destruction,by anecic and endogeic earthworms in repacked soil cores

Although the role of earthworms in soil functioning is often emphasised, many important aspects of earthworm behaviour are still poorly understood. In this study we propose a simple and cost-effective method for estimating burrow system area and continuity, as well as a new and often neglected parameter, the percentage of burrow refilling by the earthworms own casts. This novel parameter is likely to have a huge influence on the transfer properties of the burrow system. The method uses standard repacked soil cores in PVC cylinders and takes advantages of clay shrinkage and the fact that earthworms were previously shown to prefer to burrow at the PVC/soil interface. In this way, after removing the PVC cylinders off dry cores, the external section of the burrow system made by earthworms along the soil walls could be easily described. We applied this method to characterise the burrow systems of four earthworms species: two anecics (Aporrectodea caliginosa nocturna and Aporrectodea caliginosa meridionalis) and two endogeics (Aporrectodea caliginosa icaliginosa and Allolobophora chlorotica). After one month the burrow's area generated by both anecic species were much larger (about 40 cm²) than the endogeic burrow's area (about 15 cm²). A. nocturna burrow system continuity was higher than that of A. meridionalis and both anecic burrow systems were more continuous than those made by the endogeic earthworms. This was partly explained by the far larger proportion of the burrow area that was refilled with casts: approximately 40% and 50% for Al. chlorotica and A. caliginosa, respectively compared with approximately 20% for the anecic burrows. We discuss whether these estimates could be used in future models simulating the dynamics of earthworm burrow systems by taking into account both burrow creation and destruction by earthworms.     

Food quality affects production of Lumbricus terrestris (L.) under controlled environmental conditions

Birch leaves and horse manure were used to determine the effects of food quality on growth and reproduction of laboratory-reared Lumbricus terrestris. Animals grew to maturity within 6 months but attained a significantly (p < 0.001) larger adult size with manure (6.17 g) versus leaves (4.20 g). Cocoon production by recently-mated adults maintained in isolation, fed with birch leaves or horse manure, resulted in 4.53 and 3.84 cocoons ind.⁻¹ month⁻¹ respectively, with an initial hatchability of 86%, falling to zero after 18 months. Re-mating of these known individuals permitted long term monitoring of reproductive output (to 30 months). For the whole experimental period, overall hatchability of the 2010 cocoons produced was 44.4%. Median incubation time of those cocoons that hatched within accepted norms (less than 5 months at 15 °C) was 103 days and was not influenced by adult food type. A proportion (35.5%) of cocoons took in excess of 12 months to hatch. Adult mortality was minimal (25%) during the long term experiment but abnormal cocoon production was recorded after 2 years. Overall results demonstrate that food quality can have a significant influence on somatic and reproductive production of L. terrestris and these data may aid construction of production models for this earthworm in ecosystems with contrasting food quality.     

Growth of Lumbricus terrestris and Aporrectodea caliginosa in an acid forest soil, and their effects on enchytraeid populations and soil properties

A laboratory experiment was carried out to test the hypothesis that the earthworms Lumbricus terrestris and Aporrectodea caliginosa are able to maintain their populations and reproduce in the acid forest soil of a deciduous forest where no lumbricids were found in the field. The experiment was conducted in 45-l containers in which layers of mineral subsoil, humus and organic topsoil collected from the site were established. Both species survived and at least L. terrestris reproduced during the 60 weeks’ incubation. Burrows and middens of L. terrestris were recorded and quantities of litter were consumed. The presence of lumbricids increased the organic matter content of humus, reduced the acidity of the topsoil and humus layers, and suppressed the population of the enchytraeid Cognettia sphagnetorum. A dense population of Enchytraeus albidus was found in L. terrestris middens. It is concluded that edaphic factors do not explain the absence of earthworms, but isolation from cultural landscapes and lack of opportunity to colonize the site from the surroundings is the decisive factor.     

Role of the anecic earthworm Lumbricus terrestris L. in the distribution of plant residue nitrogen in a corn (Zea mays)–soil system

While the benefits of earthworms to crop production are widely acknowledged, the mechanisms involved are poorly understood. We examined the effects of an anecic earthworm (Lumbricus terrestris) on the distribution of plant residue N in a corn (Zea mays)/soil system. Soil (mixed Ap and B horizons) mesocosms (10 cm diameter, 39 cm deep) were amended with ¹⁵N-labeled corn litter, inoculated with one earthworm per mesocosm (WORM) or none (CTRL), and pre-incubated for 1, 2 or 3 weeks. Earthworms and remaining plant residues were removed and sweet corn grown in the mesocosms in a greenhouse for 3 weeks. Litter, earthworms, shoots, roots and bulk and burrow soil were analyzed for total N and ¹⁵N. Plant and earthworm biomass were also determined. Earthworms had no significant effect on the N content of shoots, roots or bulk soil. Recovery of ¹⁵N ranged from 92.6 to 101.9% in CTRL and 60.2 to 83.2% in the WORM treatment. The ¹⁵N content of bulk soil in the WORM treatment was significantly higher than in CTRL and increased with pre-incubation time. Excess at.% ¹⁵N of burrow soil was 10–100 times higher than in bulk soil. Incorporation of ¹⁵N by shoots and roots was significantly higher in the WORM treatment and increased significantly with pre-incubation time only in the WORM treatment. In WORM mesocosms pre-incubated for 3 weeks, the distribution of added ¹⁵N was 9.8% in litter, 6.5% in plant, 31.5% in soil, 12.0% in earthworms and 39.8% presumably lost as gas; in CTRL mesocosms, the values were 75.7% in litter, 3.2% in plant, 13.7% in soil and 7.4% in presumed gas losses. The activities of L. terrestris altered the distribution of plant residue N significantly, increasing the transfer of N to plants and soil and enhancing losses of N in the gas phase as pre-incubation time increased.     

Accumulation of zinc and its effects on the growth, reproduction and life cycle of Drawida willsi (Oligochaeta), a dominant earthworm in Indian crop fields

 Accumulation of Zn and its effects on the growth, reproduction and life cycle of the earthworm Drawida willsi were determined. D. willsi did not reveal any significant changes in their mass at any of the concentrations of Zn (50, 200 and 400 mg kg^–1) compared to in untreated soils. The Zn concentrations in the exposed earthworms were significantly increased, but they were able to regulate their body content of Zn within a range of 116–125 mg kg^–1 (dry wt) in 200–400 mg kg^–1 Zn-treated soil. Reproduction was significantly reduced when the Zn concentration in soil exceeded 200 mg kg^–1. The drop in reproduction at elevated concentrations of Zn apparently resulted in a delay in completion of the life cycle and a decline in the total population. Received: 9 September 1998     

Seasonal variations in growth and reproduction of the earthworm Perionyx excavatus (Oligochaeta: Megascolecidae)

 One-week-old juveniles of Perionyx excavatus were cultured in urine-free cow manure for 16 weeks in three different seasons to find out the influence of environmental factors on growth and reproduction. The biomass of worms increased gradually until the 16th week in all seasons. There was no significant (F=2.51) variation in biomass between the three seasons; however, the growth rate varied significantly (F=50.49, P<0.01). During the pre-clitellar period, the growth rate (mg/g) increased steeply (1013.17±41.33 in the monsoon season, 728.64±15.5 in winter and 463.16±22.96 in summer) until the worms attained maturity (5–8 weeks) and from then onwards gradually decreased after the commencement of cocoon production. Worms attained maturity when the mean weight of each worm ranged from 167.46 mg to 197.13 mg. The cumulative number of cocoons varied significantly (F=26.07, P<0.01) with season. Similarly, the rate of cocoon production also varied significantly (F=62.06, P<0.01). The average rate of cocoon production was 7.23, 0.99 and 0.53/worm/week in the monsoon season, winter and summer, respectively. By week 16, the ratio of weight gain to the initial weight of 1-week-old juveniles was 1 : 43.13 in the monsoon season, 1 : 30.49 in the winter and 1 : 23.69 in the summer. The maximum mass gain, growth rate, and cocoon production and earliest attainment of maturity were recorded during the monsoon season, and were attributed to the lowest range of fluctuating temperatures and high humidity compared with during winter and summer. The growth rate of worms in all seasons was inversely proportional to the number of cocoons produced. Further, the higher the rate of cocoon production, the lower the weight of individual cocoons. Received: 1 July 1997     

Genetic variation in desiccation tolerance of Dendrobaena octaedra cocoons originating from different climatic regions

The aim of this study was to examine genetic variation in desiccation tolerance in cocoons of the parthenogenetically reproducing earthworm Dendrobaena octaedra by comparing populations originating from different geographic regions (Denmark, Norway and Finland), representing large differences in precipitation and temperature. In one experiment, the tolerance of the three populations to increasing desiccation stress in the range from 100 to 91.6% relative humidity (RH) was examined, aiming to represent ecologically relevant RH values. In a second experiment, the effect of cocoon size on desiccation tolerance was investigated at 92.3% RH in the same three populations. There were highly significant differences in desiccation tolerance between populations, indicating a high genetic differentiation of this trait in D. octaedra. Cocoons from Denmark were much more sensitive (71±14% mortality at 91.6% RH) than cocoons from Norway (21±4% mortality) and Finland (4±5% mortality). Cocoons of worms from Finland and Norway were significantly larger than cocoons produced by worms from Denmark suggesting that cocoons from Denmark lost water at a higher rate when subjected to low humidity. Assuming that slow dehydration is necessary for physiologically based protection mechanisms it may be expected that desiccation tolerance is positively correlated with cocoon size. However, within each of the populations cocoon fresh weight did not have any significant impact on desiccation tolerance. When all populations were pooled there was a significant positive effect of cocoon fresh weight on desiccation tolerance, explaining about 20% of the total variation (linear regression). It seems therefore that genetic variation of desiccation tolerance in D. octaedra cocoons is related to variation in both cocoon size and other, physiologically based tolerance mechanisms.     

生物耕作对土壤理化特性、酶活性及青花菜生长和品质的影响

通过田间试验,研究生物耕作(接种蚯蚓)对土壤理化性质、酶活性及青花菜生长和产量品质的影响。试验设置了传统机械旋耕(CK)、免耕(T1)以及生物耕作(T2)3个处理。研究结果表明:(1)生物耕作能有效提高耕层不同部位的土壤养分含量和含水量,其中尤以5～20 cm土层较为明显,生物耕作处理土壤有机质、全氮、有效磷含量以及含水量依次比传统机械旋耕(CK)增长58.33%、68.93%、67.06%和16.19%;与传统机械旋耕(CK)相比,生物耕作处理大大提升了土壤过氧化氢酶、脲酶以及蔗糖酶的活性,其中脲酶、蔗糖酶与CK之间差异显著(P<0.05)。(2)生物耕作处理下青花菜的株高、开展度、叶长、叶宽等指标与CK和T1之间差异显著(P<0.05);各处理间青花菜的叶绿素相对含量差异不大,生物耕作处理增强了青花菜在各生长期的净光合速率。(3)生物耕作处理大幅提高了青花菜品质,生物耕作处理下的青花菜Vc含量是CK的1.2倍,硫代葡萄糖苷含量分别比T1处理和CK高1.05 mol·g-1和3.29 mol·g-1,差异均达到显著水平;各处理间可溶性糖含量差异不显著。因此,在传统的农业生态系统中,培育土壤有益动物的生物数量(生物耕作)可提高土壤养分和酶活性,对改善农田土壤肥力有着重要意义。     

Effects of heavy metals on the litter consumption by the earthworm Lumbricus rubellus in field soils

Aim of this study was to determine effects of heavy metals on litter consumption by the earthworm Lumbricus rubellus in National Park the “Brabantsche Biesbosch”, the Netherlands. Adult L. rubellus were collected from 12 polluted and from one unpolluted field site. Earthworms collected at the unpolluted site were kept in their native soil and in soil from each of the 12 Biesbosch sites. Earthworms collected in the Biesbosch were kept in their native soils. Non-polluted poplar (Populus sp.) litter was offered as a food source and litter consumption and earthworm biomass were determined after 54 days. Cd, Cu and Zn concentrations were determined in soil, pore water and 0.01 M CaCl₂ extracts of the soil and in earthworms. In spite of low available metal concentrations in the polluted soils, Cd, Cu and Zn concentrations in L. rubellus were increased. The litter consumption rate per biomass was positively related to internal Cd and Zn concentrations of earthworms collected from the Biesbosch and kept in native soil. A possible explanation is an increased demand for energy, needed for the regulation and detoxification of heavy metals. Litter consumption per biomass of earthworms from the reference site and kept in the polluted Biesbosch soils, was not related to any of the determined soil characteristics and metal concentrations.     

Effects of yeast on the growth and reproduction of the saprophagous millipedePolydesmus angustus (Diplopoda, Polydesmidae)

The millipedePolydesmus angustus was reared in the laboratory from hatching to maturity, reproduction and death. Two food types were used: dead leaves alone or dead leaves supplemented monthly with dry food yeast (Saccharomyces cerevisiae) at a rate not exceeding 5% of leaf dry weight. Growth, survival, adult live weight and fertility were compared between females reared on the two diets. Although the species was able to complete its life cycle on dead leaves alone, several parameters were strongly affected by the addition of yeast: growth was significantly faster, adult females became significantly larger and there was a 4.3-fold increase in fertility. Only survival was unaltered by the addition of yeast. The comparison between these laboratory results and field data on female fertility and live weight suggests that the natural diet of millipedes includes foods of higher quality than dead leaves. Possible sources of high-quality food in natural conditions are discussed.     

Effects of temperature regime on the respiratory activity of developmental stages of Lumbricus rubellus (Lumbricidae)

Rates of carbon dioxide production of cocoons, juveniles and matures of Lumbricus rubellus were measured at five constant (2°C, 5°C, 10°C, 15°C, 20°C) and three diurnally fluctuating temperature regimes (0–10°C, 5–15°C, 10–20°C) covering the whole range of temperature conditions experienced by this species in forests in Central Germany. Respiration rates of developmental stages significantly increased with temperature both under constant and fluctuating regimes. Overall, at constant temperature regimes Q₁₀ values for cocoons, juvenile and mature earthworms were 1.6, 2.7 and 2.0, respectively. At fluctuating temperature regimes Q₁₀ values were generally higher with 2.4, 3.6 and 3.5 for cocoons, juvenile and mature earthworms, respectively. At the same mean temperature respiration rates at fluctuating regimes exceeded those at constant regimes in all developmental stages.     

The influence of feeding patterns on growth and reproduction of the vermicomposting earthworm Eisenia fetida (Oligochaeta)

Summary Observations in outdoor culture beds indicated that food availability and frequency of feeding influenced worm density. In order to study the role of feeding status on the growth and reproduction of Eisenia fetida, pre-clitellate specimens were kept for 180 days under controlled conditions. A control group was regularly provided with freshly produced cattle manure while the experimental groups were fed only sporadically and at some stage even starved by removing some of the substrate. Worm growth, maturation, and cocoon production were monitored. The role of food availability was highlighted by this study. Both worm growth and cocoon production were correlated closely with the feeding pattern that was followed. This study also showed that freshly produced, urine-free, cattle manure proved an excellent food source provided it was added in such a way as to prevent the development of anaerobic conditions. Regular feeding gave high growth and reproduction rates.