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Earthworms, as the ecosystem engineers, both directly and indirectly affect the nitrogen(N) cycle. We aimed to provide a quantitative assessment of the contribution of earthworms to the terrestrial ecosystem N cycle using meta-analysis of 130 publications selected. The natural logarithm of the response ratio(lnRR) was used to indicate the effect size of earthworms on N dynamic variables. The results showed that earthworms significantly affected soil N-cycling microorganisms, including the amoA gene abundance of soil ammonia-oxidizing bacteria(AOB), and significantly promoted soil N cycle processes,including denitrification, mineralization, and plant assimilation. The effects of earthworms on the N cycle were experimental design dependent and affected by factors such as the functional group of earthworm and residue input. The presence of the anecic earthworms decreased the rates of mineralization and nitrification, and increased nitrification and denitrification responses were more pronounced in the presence of the endogeic earthworms than that of the other two functional groups of earthworms. In addition, residue input enhanced the effects of earthworms on the N cycle. The effects of earthworms on nitrous oxide(N2 O) emission increased when residues were added. These findings indicate that residue input and introducing suitable functional groups of earthworms into the field can lead to N sustainability without increasing N2 O emission. This meta-analysis also provides systematic evidence for the positive effects of earthworms on the plant N pool, N availability(soil ammonium(NH4+) content), and soil microbial biomass N content, showing the potential to alter ecosystem functions and services in relation to N cycling.  相似文献   

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There is increasing evidence that direct interactions between earthworms and seeds impact the assembly of plant communities. However, effects of earthworms of different ecological groups and their interactions on plant germination and establishment are little known. We set up a full-factorial greenhouse experiment in order to explore impacts of different ecological groups of earthworms (epigeic, endogeic and anecic) on the establishment of seedlings. The three ecological groups of earthworms affected seedling establishment in an interactive way with the effects varying in time. While anecic earthworms detrimentally affected the number of established seedlings, impacts of epigeic and endogeic species depended on the presence of earthworms belonging to other ecological groups. Impacts of anecic earthworms were more significant and consistent than those of epigeic and endogeic ones pointing to the dominance of the effect of anecic earthworms for seedling establishment. The initial positive effect of the combined presence of epigeic and endogeic earthworms compared to treatments with only endogeic and only anecic earthworms was likely due to the acceleration of seed germination and elevated nutrient availability for seedlings in earthworm casts. By contrast, reduced numbers of seedlings in presence of both epigeic and endogeic earthworms compared to the control treatment might have been due to seedling predation after the litter layer had disappeared. The results extend current knowledge on interactions between earthworms and seeds by showing that, in addition to anecic species, also epigeic and endogeic species influence seedling establishment with their effect depending on the presence of anecic species. Moreover, the results indicate that impacts of earthworms vary with the developmental stage of seedlings highlighting the importance of interactions between earthworms and seeds.  相似文献   

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Background, Aim and Scope   Background, Aims and Scope. The earthworm field test firstly was developed by the German Federal Biological Research Centre for Agriculture and Forestry (BBA) for testing of pesticides and later internationally standardised by the International Organization for Standardization (ISO) as a tool for characterizing soil quality. It is mainly used for the assessment of effects of pesticides on earthworms under field conditions. Growing experience has shown that the practical performance of the test can be improved. Main Features: In two meetings organised by the Federal Biological Research Centre for Agriculture and Forestry (Braunschweig, 2002) and later the German Federal Agency for Consumer Protection and Food Safety (Lille, 2005) an ad-hoc working group of experts from various countries and institutions discussed the practical performance and recommendations that should be taken into account in a revised draft following the current ISO guideline after the proposed revision has been approved by voting in the periodical review. Results: The outcome of the discussions and the background of the recommendations of the ad-hoc working group are briefly summarised in this paper. These are mainly methodological details in order to improve the test performance (e.g. by decreasing variability or increasing practicability). Discussion: These recommendations are proposed to be considered for the next update of the current ISO guideline. They should be seen as technical amendments still waiting for their approval by standardisation organisations. Conclusions: Although this method focuses on the assessment of effects of pesticides on earthworms, the recommendations will also be valuable for testing contaminated soils. The recommendations presented may also be a first step for splitting the current guideline into two fields of application, one for testing of chemicals and pesticides (i.e. within the scope of OECD) and another one for testing soil quality (i.e. within the scope of ISO). Perspectives: Nevertheless, there is still a major need for guidance concerning the interpretation of effects determined in such complex field tests.  相似文献   

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蚯蚓体表液的组成及其减粘脱土机理分析   总被引:16,自引:2,他引:16  
蚯蚓的体表液利于减粘脱土。本文利用现代测试仪器,结合生物化学分析技术,研究了蚯蚓体表液和泥鳅分泌液的组成;并应用土壤力学和生物化学等方面的理论,初步探讨了蚯蚓体表液减粘脱土的机理。  相似文献   

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Effects of earthworms on nitrogen mineralization   总被引:13,自引:0,他引:13  
The influence of earthworms (Lumbricus terrestris and Aporrectodea tuberculata) on the rate of net N mineralization was studied, both in soil columns with intact soil structure (partly influenced by past earthworm activity) and in columns with sieved soil. Soil columns were collected from a well drained silt loam soil, and before the experiment all earthworms present were removed. Next, either new earthworms (at the rate of five earthworms per 1200 cm3, which was only slightly higher than field numbers and biomass) were added or they were left out. At five points in time, the columns were analyzed for NH 4 + , NO 3 , and microbial biomass in separate samples from the upper and lower layers of the columns. N mineralization was estimated from these measurements. The total C and N content and the microbial biomass in the upper 5 cm of the intact soil columns was higher than in the lower layer. In the homogenized columns, the C and N content and the microbial biomass were equally divided over both layers. In all columns, the concentration of NH 4 + was small at the start of the experiment and decreased over time. No earthworm effects on extractable NH 4 + were observed. However, when earthworms were present, the concentration of NO 3 increased in both intact and homogenized cores. The microbial biomass content did not change significantly with time in any of the treatments. In both intact and homogenized soil, N mineralization increased when earthworms were present. Without earthworms, both type of cores mineralized comparable amounts of N, which indicates that mainly direct and indirect biological effects are responsible for the increase in mineralization in the presence of earthworms. The results of this study indicate that earthworm activity can result in considerable amounts of N being mineralized, up to 90 kg N ha–1 year–1, at the density used in this experiment.  相似文献   

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Relationships between anecic earthworms (Lumbricus terrestris and Aporrectodea giardi) and the collembolan species Heteromurus nitidus (Templeton, 1835), which is known to be attracted to earthworms, were investigated in an 8-week laboratory experiment. Our aims were (1) to assess whether earthworms influence the population dynamics of H. nitidus, and (2) to study pathways of influence and how earthworm effects are modified by humus forms and predators. Using microcosms with three defaunated humus forms, then provided with earthworms and predators, we intended to demonstrate that, amongst possible favourable effects of earthworms on springtail populations, earthworm activity may provide greater access and more pathways for springtails to explore soil and avoid predation. We expected that the effects of predators (centipedes) on the abundance of H. nitidus would increase from less (calcic mull) to more (moder) compact soil, and we hypothesized that earthworms would reduce predation pressure on H. nitidus by providing escape routes through increased macroporosity. Humus forms and earthworms only affected the population size of H. nitidus under high predation pressure, when collembolan numbers were higher in calcic mull than in moder, and were increased by the presence of earthworms. These results corroborate the hypothesis that earthworms, by increasing soil macroporosity, improve the escape routes for Collembola and thus evade predation. In moder humus earthworms increased the density of H. nitidus whether predators were present or not, so we cannot exclude that earthworms were also directly beneficial to H. nitidus. However, the hypothesis of a functional relationship mediated by soil macroporosity seems relevant since it was supported by differences observed when considering body size. When two size classes were distinguished within populations of H. nitidus (1) the positive effect of earthworms in moder was observed only on larger Collembola (>1 mm), (2) the density of the larger Collembola was decreased by predation only in moder and not in mull, (3) the effects of predators on the smaller individuals were not influenced by the presence of earthworms whatever the humus form, and was not decreased by the presence of earthworms. Nevertheless, factors other than macroporosity may operate as the presence of earthworms in acidic mull led to an unexplained decrease in the abundance of small-sized H. nitidus.  相似文献   

10.
We studied the effects of maize residue application on some life-cycle parameters of the earthworm Aporrectodea trapezoides in saline agricultural soils with electrical conductivity (EC) ranging from 1.58 to 7.35 dS m−1. This experiment was carried out under controlled laboratory conditions for 150 days. Results showed that soil salinity significantly affected the growth and reproduction of earthworms, decreasing survival, numbers and mean fresh weights of adults, juveniles and cocoons. Maize residue application gave a greater survival of earthworms at all salinity levels, but the differences were only significant at an EC of 7.35 dS m−1, although the mean weight of adult earthworms was significantly increased by maize residue application at all salinity levels. At an EC of 1.58 dS m−1 and 3.35 dS m−1, the application of maize residues gave significantly higher numbers of cocoons and juveniles, but in soils with 5.26 dS m−1 and 7.35 dS m−1 earthworms did not produce any cocoons over the experimental period, irrespective of maize residue application. These results indicated that maize residue application alleviated the negative effects of soil salinity on the growth and reproduction of A. trapezoides up to 3.35 dS m−1, above which maize residues only increased the growth but not on the reproduction of earthworms.  相似文献   

11.
Radish plants infested with a soil-borne plant pathogen, Fusarium oxysporum f. sp. raphani PEG-4, which is resistant to hygromycin B, were placed on the surface of a soil microcosm containing earthworms (Pheretima sp.). The earthworms ate the radish plants and scattered individual casts everywhere in the burrows. The fungal propagules were detected in the gut of the earthworms and in 26 out of 28 casts. These results suggested that the pathogen could survive passage through the alimentary canal of the earthworms and be disseminated anywhere earthworms wandered. The collected casts were incubated in a moist chamber at 28°C for 14 days, which resulted in a decline in the number of the propagules. The effects of the earthworms on the population dynamics of the pathogen were estimated. The earthworms seemed to cause a decline in total propagules of the pathogen in soil, although they expanded its distribution in soil.  相似文献   

12.
As a sensitive biological indicator, earthworms are widely used to monitor various pollutants of soil and provide an early warning for soil pollution.However, because many indices are involved in the exposure-induced oxidative stress response, practical applications of these indices are quite inconvenient.Therefore, it is appropriate to investigate the key monitoring index for use in early warning and pollution monitoring. Using Eisenia fetida as an experimental model in an indoor simulation experiment, the mathematical modelling of the effect on oxidative stress in earthworms under cadmium(Cd) stress was studied.The test lasted 40 d, with the removal of one earthworm every 10 d. The Cd~(2+)concentration gradient was set as 0, 1, 10, 20, 100, 200, 400, and 800 mg kg~(-1) dry weight. The earthworms were divided into two sections from the clitellum for the determination of total protein(TP) and peroxidase(POD), superoxide dismutase(SOD), glutathione-S-transferase(GST), glutathione peroxidase(GPX), catalase(CAT), malondialdehyde(MDA), and acetylcholinesterase(AChE)activities. Results showed that POD was the key index of oxidative stress in head tissues after 10 d of exposure, TP was the key index at 20 d, and POD became the key index again at 30 and 40 d. By contrast, in tail tissues, MDA and SOD were the key indices at an exposure time of 10 d, GPX at 20 d, CAT and TP at 30 d, and POD and MDA at 40 d. These results contribute to establishing a scientific method for ecotoxicological diagnosis and revealing the mechanism of soil Cd toxicity.  相似文献   

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Earthworms are the major component of the soil fauna in temperate agro-ecosystems. Land use and soil management are widely reported to influence earthworm populations. We report simple laboratory experiments in which earthworm survival was tested against uniaxial loads for a range of soil conditions. Across all the experimental conditions 86% of earthworms survived. While greater loads (up to 800 kPa) over longer exposure times (up to 60 s) decreased survival; even under the most severe test conditions 33% of earthworms survived. Our results suggest that decreased earthworm populations in compacted soil are not due to uniaxial loading alone, but may be the result of shearing the soil during loading or from changes to the soil properties.  相似文献   

14.
Trace metals and metalloids (TMM) in forest soils and invasive earthworms were studied at 9 sites in northern New England, USA. Essential (Cu, Mo, Ni, Zn, Se) and toxic (As, Cd, Pb, Hg, U) TMM concentrations (mg kg−1) and pools (mg m−2) were quantified for organic horizons (forest floor), mineral soils and earthworm tissues. Essential TMM tissue concentrations were greatest for mineral soil-feeding earthworm Octolasion cyaneum. Toxic TMM tissue concentrations were highest for organic horizon-feeding earthworms Dendobaena octaedra, Aporrectodea rosea and Amynthas agrestis. Most earthworm species had attained tissue concentrations of Pb, Hg and Se potentially hazardous to predators. Bioaccumulation factors were Cd > Se > Hg > Zn > Pb > U > 1.0 > Cu > As > Mo > Ni. Only Cd, Se, Hg and Zn were considered strongly bioaccumulated by earthworms because their average bioaccumulation factors were significantly greater than 1.0. Differences in bioaccumulation did not appear to be caused by soil concentrations as earthworm TMM tissue concentrations were poorly correlated with TMM soil concentrations. Instead, TMM bioaccumulation appears to be species and site dependent. The invasive A. agrestis had the greatest tissue TMM pools, due to its large body mass and high abundance at our stands. We observed that TMM tissue pools in earthworms were comparable or exceeded organic horizon TMM pools; earthworm tissue pools of Cd were up 12 times greater than in the organic horizon. Thus, exotic earthworms may represent an unaccounted portion and flux of TMM in forests of the northeastern US. Our results highlight the importance of earthworms in TMM cycling in northern forests and warrant more research into their impact across the region.  相似文献   

15.
It is well known that organic matter in the form of dung is utilised as a food source by some earthworms, but little has been reported on the preferences of earthworms for different types of dung in agricultural settings. An experiment was carried out in spring in south-eastern Australia to evaluate the effect that dung from different livestock has on the abundance of earthworms in a grassland environment. We were particularly interested to compare the responses of native Australian earthworms (Megascolecidae) with those of exotic earthworms (Lumbricidae and Acanthodrilidae). The attractiveness of dung from sheep, cattle and horses was measured by determining the abundance and biomass of the resident earthworm species under each dung type at varying times after adding the dung to the soil surface (0, 10, 20 and 30 days). The earthworm population consisted of three exotic species, Apporrectodea trapezoides, Microscolex dubius and M. phosphoreus, and two native species, Spenceriella macleayi and S. bywongensis. Both the number of days that the dung was available to earthworms and the type of dung influenced the numbers and biomass of the earthworms found beneath the dung pats. Significant interactions existed between time and dung type when all adult earthworms were considered as one group and also when adults were split into individual species. The various species responded differently to the dung, but horse dung was generally the more preferred dung type. The significance of these results is discussed in terms of the management of dung in an Australian pastoral context.  相似文献   

16.
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.  相似文献   

17.
In forest ecosystems, earthworms and wildfire are two ecological agents that cause carbon (C) stored in the forest floor to be transferred to the atmosphere as greenhouse gases, either through heterotrophic respiration (earthworms) or through periodical combustion (wildfire). For centuries, wildfire has been an important ecological driver in the boreal forests of Canada where most fire emissions to the atmosphere originate from the forest floor. In contrast, earthworms are recent invaders, having been introduced to the Canadian boreal during the 20th century. Their spread is mainly associated with anthropogenic activities. We examined stand-level effects of earthworms and wildfire on forest floor C by adapting an earthworm-C simulation model for the boreal and using it in combination with a forest C accounting model. Because the overall impact of an invasive species depends on its areal extent, we used a spatial model of earthworm spread to calculate the total predicted change in C storage at the landscape-level following earthworm invasions in northeastern Alberta. Depending on the ecological groups of earthworms modelled in stand-level simulations, the forest floor C stock was reduced by 49.7–94.3% after 125 years, although the majority of this reduction occurred 35–40 years after initiation of the invasion. Because earthworm activities reduced the amount of forest floor C available for burning, emissions from wildfire were lower in the presence of earthworms. Spatial modelling of earthworm effects within the 5,905,400 ha Alberta–Pacific Forestry Management Area projected that forest floor C stocks in the invaded stands decreased 50,875 Mg C by 2006, and 2,706,354 Mg C by 2056, compared with the same area if earthworms were not present. Loss of forest floor C averaged over the 50 year simulation was 10 g m2 yr−1; similar in magnitude to estimates for C loss in the Canadian boreal due to wildfire or harvesting. These results indicate effects of non-native earthworms on the forest floor should be included in predictions of forest ecosystem C budgets to ensure accurate attribution of emissions to heterotrophic respiration versus combustion.  相似文献   

18.
As a sensitive biological indicator, earthworms are widely used to monitor various pollutants of soil and provide an early warning for soil pollution. However, because many indices are involved in the exposure-induced oxidative stress response, practical applications of these indices are quite inconvenient. Therefore, it is appropriate to investigate the key monitoring index for use in early warning and pollution monitoring. Using Eisenia fetida as an experimental model in an indoor simulation experiment, the mathematical modelling of the effect on oxidative stress in earthworms under cadmium (Cd) stress was studied. The test lasted 40 d, with the removal of one earthworm every 10 d. The Cd2+ concentration gradient was set as 0, 1, 10, 20, 100, 200, 400, and 800 mg kg-1 dry weight. The earthworms were divided into two sections from the clitellum for the determination of total protein (TP) and peroxidase (POD), superoxide dismutase (SOD), glutathione-S-transferase (GST), glutathione peroxidase (GPX), catalase (CAT), malondialdehyde (MDA), and acetylcholinesterase (AChE) activities. Results showed that POD was the key index of oxidative stress in head tissues after 10 d of exposure, TP was the key index at 20 d, and POD became the key index again at 30 and 40 d. By contrast, in tail tissues, MDA and SOD were the key indices at an exposure time of 10 d, GPX at 20 d, CAT and TP at 30 d, and POD and MDA at 40 d. These results contribute to establishing a scientific method for ecotoxicological diagnosis and revealing the mechanism of soil Cd toxicity.  相似文献   

19.
Endogeic earthworms play an important role in mobilisation and stabilisation of carbon and nitrogen in forest and arable soils. Soil organic matter is the major food resource for endogeic earthworms, but little is known about the size and origin of the organic matter pool on which the earthworms actually live. We measured changes in body mass of juvenile endogeic earthworms, Octolasion tyrtaeum (Savigny), in soils with different C and N contents resulting from different fertiliser treatments. The soil was taken from a long-term experiment (Statischer Düngungsversuch, Bad Lauchstädt, Germany). The treatments included (1) non-fertilised soil, (2) NPK fertilised soil, (3) farmyard manure fertilised soil and (4) NPK + farmyard manure fertilised soil. The soil was incubated in microcosms with and without one juvenile O. tyrtaeum for 80 days.Earthworm biomass decreased in non-fertilised soil by 48.6%, in NPK soil by 9.4%, but increased in farmyard manure soil by 19.7% and 42.8% (soil with additional NPK application). In farmyard manure treatments the biomass of bigger individuals decreased, but in smaller individuals it increased. In NPK fertilised soil without farmyard manure only small O. tyrtaeum increased in body mass, whereas in the non-fertilised soil all individuals decreased in body mass. Generally, soil respiration correlated positively with soil carbon content. Earthworms significantly increased soil respiration and nitrogen leaching and this was most pronounced in farmyard manure treatments. Microbial activity was generally higher in farmyard manure soil indicating that farmyard manure increases labile organic matter pools in soil. Also, biomass of earthworms and microorganisms was increased in farmyard manure soil. The presence of earthworms reduced microbial biomass, suggesting that earthworms feed on microorganisms or/and that earthworms and soil microorganisms competed for similar organic matter pools in soil. The results demonstrate that NPK fertilisation only is insufficient to sustain O. tyrtaeum, whereas long-term fertilisation with farmyard manure enables survival of endogeic species due to an increased pool of utilisable soil organic matter in arable soil.  相似文献   

20.
The contribution of an earthworm species ( Amynthas vittatus ) to the increase of the nitrogen content of soil was examined. Three specimens of adult earthworms were introduced into 300 g of soil (Gray Lowland soil, silty clay) supplemented with 1% carboxymethyl cellulose in a container and incubated for 32 d at 22°C in the dark. The contents of total-N, NH4-N and NO3-N, and the population of aerobic nitrogen-fixing bacteria in soil significantly increased after incubation with the earthworms, while the natural abundance of 15N (δ15N) in soil decreased. The amount of nitrogen in the earthworms did not decrease during the incubation in the microcosm. Both acetylene reduction activity of the microcosm and incorporation of 15N to soil from atmospheric 15N2 were significantly enhanced by the introduction of the earthworms into soil, though the observed increment of nitrogen in soil was much higher than the estimated one based on the nitrogen-fixing activity. The results obtained in the present study indicated that the earthworms increased the nitrogen content of soil, presumably due to the enhancement of the nitrogen-fixing activity of the soil from the microcosm by the earthworms.  相似文献   

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