Lumbricus terrestris L. distribution within an experimental humus mosaic in a mountain spruce forest

 An experiment was designed at a mountain site to study the distribution of adult Lumbricus terrestris in relation to a small-scale mosaic of humus forms representative of different stages of a spruce forest ecosystem. Good agreement was found between distribution in the mosaic and that in the field. ANOVA tests demonstrated the strong influence of humus form on earthworm abundance when comparing a vermimull (high earthworm burrowing activity) taken from a spruce regeneration site (61.8 individuals m^–2) with a leptomoder (no earthworm burrowing activity) taken from a 60-year-old spruce stand (6.2 individuals m^–2). Other humus forms were intermediate (mean density 34.6 individuals m^–2). The same pattern was found with individual biomass, but with lower significance. Main differences observed in the experimental design were attributed to the immediate carrying density of the humus forms. A distinction was made between humus profiles built up with or without spruce cover. In the latter case (regeneration site and bilberry heath), the immediate carrying capacity indicated by the experimental approach overestimated the field density by a factor of 4. Under spruce this overestimate was even higher (approximately 10 times too high in an adult spruce stand (160 years old) and 30 times too high under moss cover). The increase in density due to experimental conditions was not determined for leptomoder humus accumulated under the actively growing spruce stand (60 years old) since the earthworm density was near zero in both cases. Relationships between humus form and earthworm populations are discussed. Received: 9 June 1997     

Changes in humus properties and collembolan communities following the replanting of beech forests with spruce

Differences of collembolan communities within the organo-mineral A layer were studied in relation to physico-chemical changes in humus at nine sites of beech forests (Fagus sylvatica L.) and first generation spruce stands (Picea abies (L.) Karst.), planted on former beech forest 30 years ago (Central Pyrenees, France). Changes in humus form were caused by the spruce plantation and occurred mainly within the fermentation horizon where acidifying litter accumulation increased the horizon depth. The recent replacement of beech by spruce induced a shift from mull towards moder humus forms, which is explained by the decreasing organic matter turnover rate. A significant decrease in the three exchangeable cations and pH under spruce was also observed. Collembolan species diversity within the A horizon was significantly lower under spruce at this early stage of the silvogenetic cycle. Differences between species composition of Collembola between the two forest stands is related to changes in environmental conditions (e.g. nutrient availability, soil porosity, soil moisture). This study shows how forest management practices are susceptible to modify biological activity within the A horizon under exotic conifer plantations.     

Earthworm surface casts affect soil erosion by runoff water and phosphorus transfer in a temperate maize crop

To test the hypothesis that earthworm surface casts contribute to soil erosion and nutrient transfers in a temperate maize crop, two rainfall experiments were set up. One was focused on the erodibility of earthworm casts, the second examined in how casts affect water runoff and nutrient transfers. Casts produced from anecic and endogeic earthworm species were both analyzed. Visual observations in the field showed no cast transport but only cast disintegration and transfers of particles. Erodibility of newly deposited casts was high and differed significantly between age groups. Cast erosion was significantly positively related to initial mass when young but not when old. The paradox is that despite a high cast abundance (25% of the area) and obvious cast erosion, amounts of sediment and nutrient losses (C, N and P) in the runoff were at least twice as high without, than in the presence of, surface casts. Earthworm activities were shown to act as a physical brake for soil erosion by (i) creating a surface roughness with the deposition of surface casts and (ii) reducing water runoff by associated enhanced water percolation. Once the breaking-down point of the physical resistance of casts was reached, all surface casts were quickly disintegrated and finally completely washed away. The amount of particulate phosphorus recovered in water runoff was 34.7 mg P m^{− 2}, while 128.5 mg P m^{− 2} was estimated to have been released from casts. The transfers were found to occur over a short-distance through successive deposition/suspension of soil particles in the water runoff.     

The heterogeneity of humus profiles and earthworm communities in a virgin beech forest

Thirty sites, encompassing a range of soil and vegetation conditions in the biological reserve of La Tillaie (Fontainebleau Forest, France) were investigated in April 1992. Beech (Fagus sylvatica L.) was the dominant tree species, with several developmental phases forming the forest patchwork. Sessile oak [Quercus petraea (Mattus.) Liebl.] was present but only as old relictual individuals. Gaps in the canopy cover were abundant, mostly caused by wind storms 2 years previously. The next most recent storm was 25 years before, resulting in distinct patches of full-grown trees. Humus profiles were classified and compared with the distribution of earthworm communities, canopy cover, and soil types. Geomorphology was responsible for the main part of the observed variation. Absence of lime in the substrate and direct contact with a sandstone stratum near the ground surface was associated with the absence of earthworms and the appearance of an OH horizon (moder humus). Elsewhere, earthworms were present and humus profiles did not display any OH horizon (mull or mull-like moder humus), but species composition was variable and strongly influenced by the thickness of the superficial sand deposit overlying limestone. On a thick (1 m or more) sandy substrate earthworm communities were dominated by epigeic species together with the anecic Lumbricus terrestris L. The species richness was higher on a shallower sandy substrate (50 cm) where lime was more accessible to tree roots and burrowing animals. The influence of the forest cycle of beech was visible in the latter case (covering most of the area), with an increase in the thickness of the OL and OF horizons and a decrease in endogeic earthworm populations during the phase of intense growth of beech. This fall in burrowing activity was apparent in gaps created by wind storms and fungal diseases within mature stands as early as 2 years after the fall of the trees.     

Effects of earthworms on soil enzyme activity in an organic residue amended rice–wheat rotation agro-ecosystem

The effect of earthworms on soil hydrolases (protease, urease, invertase, and alkaline phosphatase) and dehydrogenase activities was investigated in maize residue amended rice–wheat rotation agro-ecosystem. Experimental plots in the rotation had five treatments, i.e. incorporation or mulching of maize residues with or without added earthworms and an untreated control. The application of maize residues to soil without earthworms significantly enhanced the five soil enzyme activities compared with the control treatment during rice and wheat cultivation. The presence of earthworms further significantly enhanced protease activity in the soils with both incorporated and mulched maize residues during two cultivation seasons, but only significantly increased alkaline phosphatase activity in the soil with incorporated maize residue during the rice cultivation season. Invertase activity was significantly enhanced by the presence of earthworms in the soil with maize residue incorporation during two cultivation seasons. There were no changes in dehydrogenase activity when earthworms were present. Additionally, the five enzyme activities in earthworm casts were significant higher than those in the surrounding soil, especially dehydrogenase and invertase activities. Whatever the treatment, the values obtained for the enzyme activities in both soil and casts, except for dehydrogenase activity in earthworm casts, were significantly higher under wheat than those in rice-cultivated soil. These results indicate that the presence of earthworms strongly affected soil enzyme activities, depending on the method of organic residue application, and the enhanced enzyme activities of earthworm casts probably contributed to the surrounding soil enzyme activities.     

Earthworm distribution and abundance predicted by a process-based model

Earthworms are significant ecosystem engineers and are an important component of the diet of many vertebrates and invertebrates, so the ability to predict their distribution and abundance would have wide application in ecology, conservation and land management. Earthworm viability is known to be affected by the availability and quality of food resources, soil water conditions and temperature, but has not yet been modelled mechanistically to link effects on individuals to field population responses. Here we present a novel model capable of predicting the effects of land management and environmental conditions on the distribution and abundance of Aporrectodea caliginosa, the dominant earthworm species in agroecosystems. Our process-based approach uses individual based modelling (IBM), in which each individual has its own energy budget. Individual earthworm energy budgets follow established principles of physiological ecology and are parameterised for A. caliginosa from experimental measurements under optimal conditions. Under suboptimal conditions (e.g. food limitation, low soil temperatures and water contents) reproduction is prioritised over growth. Good model agreement to independent laboratory data on individual cocoon production and growth of body mass, under variable feeding and temperature conditions support our representation of A. caliginosa physiology through energy budgets. Our mechanistic model is able to accurately predict A. caliginosa distribution and abundance in spatially heterogeneous soil profiles representative of field study conditions. Essential here is the explicit modelling of earthworm behaviour in the soil profile. Local earthworm movement responds to a trade-off between food availability and soil water conditions, and this determines the spatiotemporal distribution of the population in the soil profile. Importantly, multiple environmental variables can be manipulated simultaneously in the model to explore earthworm population exposure and effects to combinations of stressors. Potential applications include prediction of the population-level effects of pesticides and changes in soil management e.g. conservation tillage and climate change.     

Effects of earthworm casts and compost on soil microbial activity and plant nutrient availability

Vermicomposting differs from conventional composting because the organic material is processed by the digestive systems of worms. The egested casts can be used to improve the fertility and physical characteristics of soil and potting media. In this study, the effects of earthworm casts (EW), conventional compost (CP) and NPK inorganic fertilizer (FT) amendments on N mineralization rates, microbial respiration, and microbial biomass were investigated in a laboratory incubation study. A bioassay with wheat (Triticum aestivium L.) was also conducted to assess the amendment effects on plant growth and nutrient uptake and to validate the nutrient release results from the incubation study. Both microbial respiration and biomass were significantly greater in the CP treatment compared to EW treatment for the initial 35 days of incubation followed by similar respiration rates and biomass to the end of the study at 70 days of incubation. Soil NO₃⁻ increased rapidly in the EW and CP treatments in the initial 30 days of incubation, attaining 290 and 400 mg N kg⁻¹ soil, respectively. Nitrate in the EW treatment then declined to 120 mg N kg⁻¹ soil by day 70, while nitrate in the CP treatment remained high. While ammonium levels decreased in the CP treatment as nitrate level increased with increasing incubation time, a low level of ammonium was maintained in the EW treatment throughout the incubation. The wheat bioassay study included two additional cast treatments (EW-N and EW2) to have treatments with higher levels of N input. Plants grown with CP or FT treatment had a lower shoot biomass and higher shoot N content than in EW-N and EW-2 treatments, and also showed symptoms of salinity stress. Ionic strength and other salinity indicators in the earthworm cast treatments were much lower than in the CP treatment, indicating a lower risk of salinity stress in casts than in compost. All cast and compost amendments significantly increased wheat P and K uptake compared to either the non-amended control or the mineral fertilizer treatment. The results show that casts are an efficient source of plant nutrients and that they are less likely to produce salinity stress in container as compared to compost and synthetic fertilizers.     

腐殖酸对汞的络合稳定特性及其环境学意义

对3种腐殖酸与Hg的络合稳定特性研究结果表明,腐殖酸与Hg的结合存在松结态与紧结态2种形态,其中富里酸主要以松结态为主,灰色胡敏酸与棕色胡敏酸则以紧结态占绝对优势。3种腐殖酸中富里酸对Hg的络合容量最高,但络合强度最低,其络合汞的环境活性将较高;灰色胡敏酸对Hg的络合容量最低,但络合稳定性最高,故所结合Hg的环境活性最弱。但受容量因素制约,灰色胡敏酸对Hg的络合稳定性将因Hg/灰色胡敏酸相对比例的升高而急剧下降。棕色胡敏酸对Hg络合特性介于富里酸与灰色胡敏酸之间。     

Influence of cropping intensity on the production and properties of earthworm casts in a leucaena alley cropping system

The effect of cropping intensity on surface cast production by earthworms was studied more than 4 years in an alley cropping system. The plots were cultivated to maize–cassava intercrop every year (100% cropping intensity), after 2 years of fallow (33% cropping intensity) and after 3 years of fallow (25% cropping intensity). Cast production was assessed by the continuous sampling technique proposed by Hauser and Asawalam (Z Pflanzenernahr Bodenkd 161:23–30, 1998). Results showed that cropping intensity enhanced cast production in the first year, did not affect it in the second and third years and decreased cast production in the fourth year. Amount of casts deposited within the alleys was highest under the hedgerow and decreased towards the middle of the alley. The concentrations of organic matter and plant nutrients were significantly higher in worm casts than in soil, irrespective of the cropping intensity and position of sample collection within the interrow space. Continuous cropping up to 6 years significantly reduced the concentrations of organic carbon, total nitrogen, available phosphorus, and exchangeable calcium, potassium and magnesium in the top 0–5 cm of the soil, irrespective of the distance from the hedgerow. Regardless of the intensity of cropping, there is a strong linear association between the quantity of casts and the amount of organic carbon, and other nutrients in casts.     

Earthworm activities and the soil system

Summary Earthworms find in soil the energy, nutrient resources, water and buffered climatic conditions that they need. According to the food resource they exploit and the general environmental conditions, earthworms can be grouped into different functional categories which differ essentially in morphology, size, pigmentation, distribution in the soil profile, ability to dig galleries and produce surface casts, demographic profiles and relationships with the soil microflora. Soil characteristics are both the determinant and the consequence of earthworm activities, since these animals greatly influence the functioning of the soil system. When present, they build and maintain the soil structure and take an active part in energy and nutrient cycling through the selective activation of both mineralization and humification processes. By their physical activities and resultant chemical effects, earthworms promote short and rapid cycles of nutrients and assimilable carbohydrates. Thus earthworms represent a key component in the biological strategies of nutrient cycling in soils and the structure of their communities gives a clear indication of the type of soil system that they inhabit.     

Earthworms change the abundance and community structure of nematodes and protozoa in a maize residue amended rice-wheat rotation agro-ecosystem

The influence of earthworms on nematodes and protozoan communities was determined during the wheat phase of a six year rice-wheat rotation agro-ecosystem. Experimental plots in the rotation had five treatments, i.e. incorporation or mulching of maize residues with or without added earthworms and a control. The addition of maize residues to soil strongly affected the abundance and community structure of nematodes and protozoa in the absence of earthworms. The presence of earthworms gave significantly lower total nematode numbers at all soil depths following maize residue incorporation than the same treatment without earthworms, and also gave lower (although not significantly) total nematode numbers in the upper soil layer following maize residue mulching than the same treatment without earthworms. This was mainly due to a significant decrease in bacterial-feeding nematode numbers. Earthworms also strongly affected the distribution of the number of total nematodes and two trophic groups (bacterial and plant feeders) with soil depth. In the presence of earthworms, total protozoan and flagellate numbers significantly increased at all soil depths following both incorporation and mulching of maize residues, while numbers of amoebae increased only when maize residues were mulched. Additionally, in earthworm casts total nematode numbers (mainly bacterial and fungal feeders) were significantly higher, whereas total protozoa numbers (mainly flagellates and amoebae) were significantly lower than that in soil from 0 to 5 cm layer.These results indicated that earthworm activity could affect the abundance and community structure of microfauna, and change their distribution between soil layers and cast material, depending on the mode of application of organic residues.     

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.     

蚯蚓纤溶酶组分及其基因的多态性

生化制备证明,蚯蚓纤溶酶为一组含有10个以上同工酶组分的混合酶。为了研究这些组分的DNA和蛋白质序列的异同,本文通过对数据库中已报道的28条蚯蚓纤溶酶基因按相似性进行归类,将它们分为4组（基因型）。同一组中的序列具有共同特征,即保守的N-末端、C-末端和相同的结构域,而且这些结构域在序列中分布的位置也相同;但它们之间在中间部分存在明显差异,这些差异说明了基因型中存在多态性。这种多态性可能是它们在体内溶栓的药理药效作用存在差异的结构基础。     

Earthworm activity and soil structure changes due to organic enrichments in vineyard systems

 The effect of organic enrichment on earthworm activity and soil structure was studied in two French vineyards, by comparing control and test plots. In each vineyard the organic matter quantitatively increased the abundance and biomass of the earthworm community. These increases were associated with a higher level of species diversity and a higher evenness corresponding to the development of endogeic community. These earthworm community changes were associated with an increase in granular bioturbated areas and in macroporosity in the top soil layer. The micromorphological approach incorporated an original process of image analysis which appeared to be an appropriate method for characterizing pore morphology in this study. The pores when characterized by their size and shape could be related to ecological groups and growth stages of earthworms. Received: 4 August 1997     

蚯蚓黏液对黑麦草幼苗生长及其对菲吸收的影响

以50 mg/kg菲人工污染黄棕壤为供试土壤,设置添加蚯蚓黏液处理和不添加蚯蚓黏液对照（(CK）),研究蚯蚓黏液对黑麦草幼苗生长及吸收菲的影响,添加蚯蚓黏液处理根据蚯蚓黏液浓度设置4个水平,分别为蚯蚓黏液原液浓度的2/8,3/8,1/2和1倍。结果显示：①蚯蚓黏液促进了黑麦草幼苗的生长,根系鲜重比对照增加了33.6% ~ 146%,地上部和整株生物量鲜重比对照增加了12.4% ~ 35.8% 和34.5% ~ 59.6%;②显著降低了根部菲的浓度,约23.0% ~ 49.0%,促进了黑麦草对菲的吸收,使吸收量比对照增加了约62.8%;③降低了黑麦草根系对菲的富集系数,与对照相比降低约33.7% ~ 52.6%,提高了菲在黑麦草体内的传输系数,约54.0% ~ 102.0%。     

木质素与蚯蚓对黑麦草生物量及土壤微生物群落的影响

采集农田土壤,设置添加木质素和蚯蚓不同组合处理的黑麦草盆栽试验,在测定植物生物量的基础上,结合定量PCR、高通量测序等方法,研究木质素和蚯蚓及其联合作用对土壤细菌、真菌群落的影响.结果显示:木质素显著降低了黑麦草的生物量,而蚯蚓能够缓解木质素的抑制作用,促进黑麦草生长.木质素明显抑制了土壤细菌、真菌数量,并改变群落结构...     

Fine-scale spatial and temporal variation in earthworm surface casting activity in agroforestry fields, western Honduras

Quantification of the spatial and temporal distribution patterns of soil fauna is a relatively new area of research, and has been proposed as the key to understanding the high diversity typical of soil fauna communities. Field research on the relationships among the spatial distribution patterns of trees, litter and earthworm surface casting was carried out in two agroforestry fields in a rugged area of western Honduras. Grid-based sampling at a scale of 2-20 m was employed to determine whether any spatial relationships existed among these variables at this fine scale. Each field was sampled twice at either 2 or 3 week intervals, to determine the short-term stability of spatial relationships. The spatial distribution of litter showed a strong pattern of aggregation, whereas earthworm cast distribution did not exhibit strong spatial autocorrelation. However, the spatial distribution patterns of each of these variables were well explained by the spatial arrangement of trees in both sites. Fitted model cross-semivariograms explained between 70% and 90% of the total variation in cross-semivariance between tree density and litter cover, and between tree density and earthworm cast weight. The results of the study suggest that farmers may be able to manipulate populations of earthworms indirectly by managing the spatial arrangement of trees within their crop fields. Planning the distribution of trees could allow farmers to create ‘patches’ of organic resources within fields, while minimising the negative effects of trees on crop growth due to competition for light, water and nutrients. Over the short time scale of the study, unusually heavy rainfall led to substantial changes in spatial distribution patterns of earthworm cast activity and litter cover, which may otherwise not have occurred. This result emphasises the need to take into account short-term temporal change during ecological studies at fine spatial scales.     

Morphological characterisation of humus forms in recent coastal dune ecosystems in Belgium and northern France

The morphology of humus profiles, developed on recent sandy coastal dune ecosystems, is studied on macro- and microscale. Humus and soil profiles were described, sampled and analysed. Undisturbed, oriented samples were taken from the surface horizons for micromorphological study. The soils are situated in the National Biological Reserve of the Opal Coast, Merlimont, France and near De Haan, Belgium. On both sites, the parent material consists of aeolian calcareous sand. Surface horizons, however, are decarbonated due to leaching processes.In Merlimont, the sequence consists of two profiles, one under Pinus nigra ssp. laricio and one under Populus nigra ssp. nigra, characterised by a Resimor¹/Dysmoder^{, 2} and Rhizomull^{, 1}/Mesomull ^{, 2} humus type, respectively. At the De Haan site, three profiles under Quercus robur, Populus ssp. and P. nigra var. austriaca are investigated. The humus types are respectively Mullmoder^{, 1} to Mull/Dysmull ^{, 2}, Mull ^{, 1}/ Oligomull ^{, 2}, Mormoder ^{, 1}/Dysmoder ^{, 2}.Differences among the humus profiles are related to presence/absence among the L-, F-, H- and A-horizons, structure, rooting, presence of hyphae, sclerotia and excrements.Macromorphologically, the difference between humus profiles under coniferous and deciduous forest/grassland is evident by thicker L-, F and/or H-horizons under coniferous trees. The microstructure of the L- and F-horizons shows a loose packing of subhorizontally oriented needles. The microstructure of the H-horizons varies from intergrain microaggregate to locally weak crumb. The microstructure of the surface mineral horizon of all profiles is mainly intergrain microaggregate, sometimes single grain and exceptionally bridged grain. Intimate mixing and complexation of organic matter to the mineral fraction into aggregates is very limited in the A-horizons. All A-horizons, irrespective of the humus type, are to some degree characterised by a “pepper and salt” appearance. The horizon symbol “H+E” is introduced to label this morphology.The Mull humus forms under deciduous trees are indicative of rather intense mesofauna activities. The absence of a crumb structure in the A-horizon of these soils could be due to the fast disintegration of earthworm casts in these sandy soils with no clay or silt fraction.