Soil partitioning effect of an earthworm burrow system

Summary Through the simulation of an earthworm burrow system and the calculation of the shortest pathway from the bottom of the system to the surface, using a succession of burrows, it is shown that the borrow system leads to a partition of soil space. The characteristics of the resulting heterogeneity, the size of this partitioning and the length of the shortest pathways are discussed with regard to the functional relationship between the burrow system and the soil.Dedicated to the late Prof. Dr. M. S. Ghilarov     

Effects of metal pollution on soil macroinvertebrate burrow systems

A reduction in the numbers of macroinvertebrates present in soil may have a negative effect on soil structure, infiltration rates, and gas exchanges. Soil pollution by metal is known to have a detrimental effect on soil macrofauna. The aim of the present study was to evaluate (1) direct and indirect effects of soil pollution on soil macroinvertebrate bioturbation and (2) effects of the two macroinvertebrate communities found in a polluted and a nonpolluted area (one supposed sensitive, the other tolerant to metals) on burrow systems parameters. Macroinvertebrate porosity was studied using X-ray tomography. Three-dimensional reconstructions and characterisation of the burrow system were obtained using image analysis. Results showed that metal pollution principally affected the spatial distribution of macropores (more macropores were found near the soil surface) and the shape of the burrow system (branching rate was higher in the polluted soil), whereas soil macroinvertebrate composition principally affects burrow density parameters (the number of burrows was higher for the sensitive macroinvertebrate community).     

Burrow systems of endogeic earthworms: Effects of earthworm abundance and consequences for soil water infiltration

By creating burrows, earthworms influence the transfer properties of soils. The effects of endogeic species on soil transfer properties, however, are not yet well understood because these earthworms generally create burrows that are refilled by casts and have no preferential vertical orientation. Thirty soil cores were incubated for various periods (1–3 or 4 weeks) at different earthworm densities (70, 210, 345 or 480 individuals m⁻²). The cores were then scanned using X-ray tomography and the burrow systems were characterised by measuring the total burrow volume, bioturbation volume (refilled burrows and lateral compaction around the burrows), the number of branches, tortuosity and continuity (assessed by computing the number of burrows with a vertical extension greater than 15, 20 and 25% of the core). We also computed the mean geodesic distance, i.e. the mean distance from the bottom to the top of the core assuming that distances inside burrows are null. Rainfall simulations were carried out on 17 cores chosen to encompass the variations observed in the burrow systems. The water transfer efficiency of each core was estimated by measuring two parameters: breakthrough volume and the percentage of water transmitted after 1 h of rain. Burrow and bioturbation volume increased significantly and steadily with time and earthworm density. We estimated that on average Allolobophora chlorotica burrowed 22 cm per week. All other burrow system characteristics also increased with time and earthworm density except the mean geodesic distance, which decreased significantly. This suggests that intraspecific interactions had no significant effect on burrow system geometry. Univariate PLS regressions were used to understand which burrow system characteristics had the strongest influence on water transfer. These regressions showed that the mean geodesic distance was the most important parameter. This means that in addition to individual burrow characteristics, the spatial arrangement of the whole burrow system also had a major effect on transfer properties.     

Impact of soil compaction on earthworm burrow systems using X-ray computed tomography: preliminary study

This study is a first approach of the impact of soil compaction due to trafficking by machinery on earthworm burrow systems. To this end, two experiments were established. In the first one, microcosms were incubated in the laboratory for 70 d with Lumbricus terrestris or Aporrectodea giardi. In the second experiment, soil cores were excavated from a sugar beet field mainly colonised by L. terrestris and Aporrectodea caliginosa. The cores were then artificially compacted at 0.12 MPa or 0.25 MPa (which corresponds to the compaction due to trafficking by machinery in the field) or remained non-compacted. The whole cores were submitted to an X-ray computed tomography scan. This method allowed to compare the characteristics of the entire burrow system (total and mean burrowed length, mean length and number of burrows) and of the burrows themselves (number, area and roundness of pores constituting the burrows) in the compacted and non-compacted cores. The results showed that soil compaction contributes to close numerous pores, reduce mean length of burrows and increase the number of fragmented burrows. We concluded that soil compaction affects to a large extent the functionality of burrow systems by fragmenting them and affecting their continuity. This impact increases with the intensity of compaction. The applied soil compaction mainly affected the burrows located in the upper part of the soil. From a general point of view and especially for the upper part of the cores, horizontal burrows seemed to be more affected by compaction than vertical or oblique ones. Globally, the effects of soil compaction were especially observed at the greatest applied pressure (0.25 MPa). At 0.12 MPa, the damage to the burrows was less obvious.     

Sublethal effects of imidacloprid on the burrowing behaviour of two earthworm species: Modifications of the 3D burrow systems in artificial cores and consequences on gas diffusion in soil

Earthworms have been termed ‘ecosystem engineers’ (sensu [Jones, C.G., Lawton, J.H., Shachak, M., 1994. Organisms as ecosysem engineers. Oikos 69, 373-386.]) because of the important roles they play in the soil. As a consequence, it is assumed that if earthworms change their behaviour following exposure to pesticides or pollutants this could have a drastic impact on soil functioning. To test this assumption under laboratory conditions, we studied the burrow systems made by two earthworm species (the anecic Aporrectodea nocturna and the endogeic Allolobophora icterica) in artificial soil cores containing imidacloprid, a widely used neonicotinoid insecticide. After 1-month incubation period, the macropores created in the soil core were analyzed by tomography. In order to further characterize transfer properties associated with burrow systems gas diffusion measurements were also carried out. The burrow systems made by the two earthworm species were very different: A. nocturna made more continuous, less branched, more vertical and wider burrows than A. icterica. Some changes to A. nocturna burrow systems were observed after exposure to imidacloprid (they made a smaller burrow system and burrows were more narrow), but only at the highest concentration of imidacloprid used (0.5 mg kg⁻¹). A. icterica worms were more sensitive to imidacloprid and many differences in their burrow systems (length, sinuosity, branching rate and number of burrows) were observed at both concentrations tested (0.1 and 0. 5 mg kg⁻¹). As a consequence, the continuity of the burrow systems made by both species was altered following imidacloprid treatment. Gas diffusion through the A. nocturna soil cores was reduced but no difference in gas diffusion was observed in the A. icterica soil cores.     

Characterisation of the three-dimensional structure of earthworm burrow systems using image analysis and mathematical morphology

The aim of this work was to exemplify the specific contribution of both two- and three-dimensional (3D) X-ray computed tomography to characterise earthworm burrow systems. To achieve this purpose we used 3D mathematical morphology operators to characterise burrow systems resulting from the activity of an anecic ( Aporrectodea nocturna), and an endogeic species ( Allolobophora chlorotica), when both species were introduced either separately or together into artificial soil cores. Images of these soil cores were obtained using a medical X-ray tomography scanner. Three-dimensional reconstructions of burrow systems were obtained using a specifically developed segmentation algorithm. To study the differences between burrow systems, a set of classical tools of mathematical morphology (granulometries) were used. So-called granulometries based on different structuring elements clearly separated the different burrow systems. They enabled us to show that burrows made by the anecic species were fatter, longer, more vertical, more continuous but less sinuous than burrows of the endogeic species. The granulometry transform of the soil matrix showed that burrows made by A. nocturna were more evenly distributed than those of A. chlorotica. Although a good discrimination was possible when only one species was introduced into the soil cores, it was not possible to separate burrows of the two species from each other in cases where species were introduced into the same soil core. This limitation, partly due to the insufficient spatial resolution of the medical scanner, precluded the use of the morphological operators to study putative interactions between the two species     

Evolution of burrow systems after the accidental introduction of a new earthworm species into a Swiss pre-alpine meadow

 The unintentional introduction of a new earthworm species (Aporrectodea nocturna) into a Swiss pre-alpine meadow resulted in a great increase in earthworm density in the newly colonized area (386 m^–2) compared with the density observed in the natural area (273 m^–2) where an earthworm community was already present. To investigate the impact of this introduction on the burrow systems, eight soil cores (length 25 cm, diameter 16 cm) were taken (four in the colonized area and four in the natural area) and analysed with computer-assisted X-ray tomography. The resulting images were processed to obtain the 3D-skeleton reconstructions of the earthworm burrow systems. Due to high variability in these burrow systems, only slight differences were observed between the two areas. The total burrow length and the mean burrow lengths tended to be greater in the colonized area. Moreover, the distribution of pore numbers with depth showed different patterns with a maximum for depths between 10 cm and 15 cm in the colonized area and a maximum for depths between 20 cm and 25 cm in the natural area. These differences may have been related to: (1) the particular behaviour of A. nocturna, which was observed to cast at the surface in this site, and (2) the predominance of juvenile earthworms around the colonization front. These differences were sufficient to create significant effects on the continuity of the burrow systems (assessed by the number of different pathways between virtual horizontal planes) for the two areas. The colonized area was characterized by a greater pore continuity, which could have resulted in enhanced transfer properties. Received: 2 July 1999     

3D skeleton reconstructions of natural earthworm burrow systems using CAT scan images of soil cores

 Four soil cores (length, 20 cm; diameter, 16 cm) were sampled in a Swiss pre-Alpine meadow with high earthworm abundance (>400 individuals/m²); two cores were taken in October 1993 and the other two cores in April 1994. The cores were described using computer assisted tomography which gives a series of section images every 3 mm. A method for reconstructing the three-dimensional (3D) skeleton of the earthworm burrow system is presented and discussed. This method provides an image of the structural organisation of the burrow system and was found to be adequately sensitive for use in ecological and functional studies. The seasonal variation of these 3D skeletons was investigated using two approaches, i.e. the analysis of: (1) global burrow system characteristics, and (2) individual burrow characteristics. At the scale of the global burrow system no difference was found between seasons (same number of burrows and same total burrow length) except for the vertical segment distribution, which was homogeneous in spring and decreased with depth in the fall. The study of individual burrow characteristics revealed that burrows tended to be more vertical in spring and that their branching intensity was higher in this season. Received: 10 June 1997     

Assessment of earthworm contribution to soil hydrology: a laboratory method to measure water diffusion through burrow walls

The capacity for water diffusion in burrow walls (i.e. the coefficient of sorptivity) either burrowed by Lumbricus terrestris (T-Worm) or artificially created (T-Artificial) was studied through an experimental design in a 2D terrarium. In addition, the soil density of earthworm casts, burrow walls (0–3 mm around the burrow) and the surrounding soil (>3 mm) were measured using the method of petroleum immersion. This study demonstrated that the quantity of water which transits through burrows of L. terrestris in the soil matrix was lower than that transited through soil fractures, due to a reduction of soil porosity in burrow walls (compaction: cast > worms burrow walls > surrounding soil > artificial burrow walls). Earthworm behaviour, in particular burrow reuse with associated cast pressing on walls, could explain the larger burrow wall compaction in earthworm burrows. If water diffusion was lower through the compacted burrows, burrow reuse by the worms makes them more stable (worms would maintain the structure over years) than unused burrows. The present experimental design could be used to test and measure the specific differences between earthworm species in their contributions to water diffusion. Probably, these contributions depend on the presumed related-species behaviours which would determine the degree of burrow wall compaction.     

铅胁迫下小花南芥与玉米间作对根系分泌物 有机酸的影响

为了揭示Pb胁迫对间作和单作的超累积植物和作物根系分泌低分子有机酸的影响,研究设置400 mg·L?1Pb胁迫,采用水培曝气法试验,以玉米和小花南芥单作为对照处理,研究Pb胁迫下玉米和小花南芥间作对植物根系形态、根系分泌有机酸及Pb吸收的影响。结果表明:与单作相比,间作小花南芥情况下,玉米根系分泌物检测到乳酸;玉米分根条数、根表面积和根密度与单作相比分别增加60%、15%和42%,地下部和地上部干重生物量分别增加108%和75%,玉米地下部Pb含量下降44%;与单作相比,间作玉米条件下,小花南芥根系分泌物检测到乙酸和乳酸,小花南芥根系分泌物量与单作相比增加103%~1 700%,小花南芥地下部和地上部Pb累积量分别比单作增加49%和75%,转运系数增加22%。相关分析结果表明,单作小花南芥只有地上部Pb累积量与草酸显著相关,而间作小花南芥地下部和地上部Pb累积量与草酸、柠檬酸和苹果酸显著相关。研究表明超富集植物小花南芥与玉米间作体系,根系分泌的有机酸改变了Pb在小花南芥和玉米体内的累积特征,促进超累积植物小花南芥累积Pb,减少农作物玉米植株体内Pb含量。Pb胁迫下超累积植物小花南芥与玉米间作是一种可行的修复模式。     

The importance of thinking big: Large-scale prey conservation drives black-footed ferret reintroduction success

Objective evaluations of wildlife reintroductions are vital for increasing the success of future efforts to re-establish endangered species. Attempts to reintroduce one of the most endangered mammals in North America, the black-footed ferret (Mustela nigripes), have been ongoing for 18 years with no quantitative assessment of factors related to reintroduction success. We examined relationships between ferret reintroduction success and factors associated with disease outbreaks, release strategies, and the distribution and abundance of their primary prey, prairie dogs (Cynomys sp.), at 11 reintroduction sites. The most important factor related to ferret reintroduction success was a cumulative metric incorporating both size of the area occupied by prairie dogs and density of prairie dog burrows within that area. Each of the four successful sites had prairie dog populations that occupied an area of at least 4300 ha. No sites with <4300 ha of prairie dogs were successful in maintaining ?30 adult individual ferrets over multiple years without augmentation even if they had a high prairie dog burrow density. The overarching importance of the availability of high-quality habitat suggests managers should prioritize actions that maintain and enhance the availability of large areas with high prairie dog burrow density, which are becoming increasingly rare due to anthropogenic impacts and disease outbreaks.     

Characterization of Portuguese populations of <Emphasis Type="Italic">Vitis vinifera</Emphasis> L. ssp. <Emphasis Type="Italic">sylvestris</Emphasis> (Gmelin) Hegi

Wild populations of Vitis vinifera L.␣have been located in Portugal. Morphological characterization was carried out in three populations located in Alcácer do Sal, Castelo Branco, and Montemor-o-Novo, and then compared using multivariate discriminant analysis. These populations were from three different hydrological basins, therefore cross-pollination was not possible. It was verified that in each population all plants were different. The data suggest that the frequency of female and male plants is rather variable in wild populations. The morphology of the adult leaf, from the Alcácer do Sal population, had particular features when compared with Castelo Branco and Montemor-o-Novo populations, which were more homogeneous. The length of teeth compared with width at the end of the base, and the density of prostrate hairs between and on main veins (lower side) were the variables which allowed the best discrimination among populations.     

旱地土壤硝态氮的产生、淋洗迁移及调控措施

如何在保证作物高产的前提下,提高氮肥利用率、减少氮肥损失及其对环境的影响已经成为一个世界性研究课题。硝态氮淋洗是旱地土壤氮素损失去向的重要途径之一,是导致区域性地下水硝酸盐污染的重要原因,已成为全球集约农区重大农业和生态环境问题。明确不同农作系统土壤硝态氮的产生、淋洗过程及影响因素,并进一步提出阻控措施,具有重要的农学和环境意义。在这一方面国内外学者已经开展了长期、大量的研究,并积累了丰富的资料。本文简要综述了旱地土壤硝化作用的微生物驱动过程,比较了当前最为常用的田间原位监测硝态氮淋洗方法的优缺点,分析了影响硝态氮累积和淋洗的各种因素,总结了相应的调控措施,并对今后的研究工作进行展望:加强旱地土壤硝化作用的关键微生物过程、机理以及相关驱动因子调控作用的研究,有助于增加对土壤氮素循环的认识,而这些认识是旱地农作系统进行氮素优化管理和制定硝态氮淋洗阻控技术的重要基础;继续寻找和建立一种扰动更少、维护简单、样品污染更低、尤其适合长期监测的田间原位研究方法,对于准确地揭示旱地土壤硝态氮的淋洗规律和实时通量尤为重要;系统地开展不同旱地农作系统硝态氮淋洗的综合性研究,阐明影响硝态氮累积和淋洗的主要因素,并提出针对性的阻控措施,更符合田间实际,对于降低区域硝态氮淋洗风险意义重大。     

Characterization of the burrow system of the earthworms Lumbricus terrestris and Aporrectodea giardi using X-ray computed tomography and image analysis

 The burrow systems of two earthworm species (Lumbricus terrestris and Aporrectodea giardi) were studied in artificially packed soil columns placed in controlled conditions in the laboratory. At the end of the incubation, which lasted 246 days, the burrow systems were characterized on the undisturbed columns using X-ray computed tomography. This method provided a set of digitized images corresponding to horizontal 3-mm-thick sections. The following parameters were measured using image analysis on each section: number of biopores, their volume, and their individual orientation calculated using an elliptical model of the earthworm channel. The profiles of these parameters through the columns showed that the burrow systems of L. terrestris and A. giardi, which are both anecic species, were very different in terms of total volume, number of burrows, burrow orientation and extension with soil depth. These results led us to conclude that the burrow system of L. terrestris can be considered as a permanent structure whereas that of A. giardi is closer to the burrow system of endogeic species. Received: 12 June 1998     

稻蟹共作对稻田水体底栖动物多样性的影响

稻蟹共作是一种生态型种养新模式,而底栖动物是该复合生态系统的重要组成部分。为研究稻蟹共作对稻田底栖动物多样性的影响,在水稻分蘖期、拔节期、扬花期和成熟期,采集不同放养密度的稻蟹共作稻田(仔蟹放养密度10 ind.m 2,低密度,LD;仔蟹放养密度30 ind.m 2,高密度,HD)和常规稻田(不放蟹,常规栽培,CK)水体的底栖动物,分析不同稻田生态系统底栖动物的种类构成、密度和多样性。结果表明:水稻分蘖期,3种放蟹密度处理稻田底栖动物的种类数、密度、多样性指数和Pielou均匀度指数均无显著差异(P>0.05);在拔节期和扬花期,养蟹稻田底栖动物的种类数和密度均小于常规稻田,且与放蟹密度呈反比;常规稻田底栖动物的Shannon-Wiener多样性指数在拔节期和扬花期均显著高于高密度养蟹稻田(P<0.05),而只在拔节期显著高于低密度养蟹稻田(P<0.05),且Pielou均匀度指数也在拔节期显著高于低密度养蟹稻田(P<0.05);成熟期,3种处理底栖动物的种类数和密度均达到最低点,Shannon-Wiener多样性指数无显著差异(P>0.05),高密度养蟹稻田的Pielou均匀度指数显著高于常规稻田(P<0.05)。研究结果可为稻蟹共作稻田底栖动物的多样性保护和稻蟹共作技术的可持续发展提供理论指导,促进稻蟹共作技术的发展和推广。     

巴丹吉林沙漠南缘近流沙区白刺灌丛沙堆形态特征与空间分布

白刺灌丛沙堆是风沙塑造的荒漠景观,研究白刺灌丛沙堆形态特征和分布格局对沙源的响应,可为进一步研究其形成演化提供依据。研究通过野外调查测定,对巴丹吉林沙漠南缘近流沙区白刺灌丛沙堆集中分布区的沙堆形态特征及其空间分布进行了研究。结果表明:雏形阶段和发育阶段的沙堆分别占沙堆总数的24.0%和42.2%。该区域相邻沙堆平均间距仅为9.8 m,沿主导风向,相邻沙堆的间距呈现由大快速减小,维持小间距一段距离后,再缓慢增大的分布格局,密度变化与此相反。受高沙源供给及由此带来的强蚀积转换的影响,白刺灌丛沙堆稳定化进程减缓,各项形态特征值均较小。相关性分析表明,白刺灌丛沙堆的高度、长轴长、短轴长、迎风坡长、背风坡长和底面积间均呈显著相关(P0.01)。研究结果说明,在丰富沙源供给条件下,研究区白刺灌丛沙堆处于发育的中早期阶段,在景观上表现出密度大而体型小的特点。     

干旱胁迫对花生生育中后期根系生长特征的影响

花生是较耐旱的经济和油料作物, 长期少雨或季节性干旱是限制花生产量提高的重要环境因子, 也是花生收获前黄曲霉素感染的重要因素。根系是植物吸水的主要器官, 不同土壤水分状况下植物的根系构型可能会表现出显著差异, 进而影响植物根系吸收养分和水分的能力。研究不同土壤水分状况下花生根系形态的发育特征与抗旱性的关系对进一步理解花生的水分吸收、运输、利用和散失机制以及培育抗旱性花生具有非常重要的作用。为明确不同抗旱性花生品种的根系形态发育特征, 探讨其根系形态发育特征对不同土壤水分状况的响应机制, 在防雨棚旱池内进行土柱栽培试验, 研究抗旱型花生品种"花育22号"和干旱敏感型花生品种"花育23号"生育中后期根系生长特征及其对干旱胁迫的响应。设置正常供水和中度干旱胁迫(分别控制土壤含水量为田间持水量的80%~85%和45%~50%)2个水分处理, 分别在花针期、结荚期和饱果期进行取样,根长、根表面积和体积扫描后通过WinRhizo Pro Vision 5.0a程序进行分析; 收获时测定产量和抗旱系数(干旱胁迫处理与正常供水处理下产量之比)。结果表明, "花育22号"具有较高的产量和抗旱系数, "花育23号"对干旱胁迫的适应性小于"花育22号"。抗旱型品种"花育22号"具有较大的根系生物量、总根长和根系表面积, 且深层土壤内根系表面积和体积大于"花育23号"。与正常供水处理相比, 干旱胁迫显著降低2个品种花针期的根系总根长、根系总表面积和总体积, 对结荚期和饱果期根系性状无显著影响; 干旱胁迫增加2个品种生育中后期40 cm以下土层内的根长密度分布比例、根系表面积和体积, 但"花育23号"各根系性状增加幅度小于"花育22号"。干旱胁迫处理下20~40 cm和40 cm以下土层内根系表面积和体积分别与总根长、总表面积和总体积呈显著或极显著正相关, 而正常供水处理下0~20 cm土层内根系表面积和体积与整体根系性状表现极显著正相关。总体而言, 具有较大根系和深层土壤内较多的根系分布是抗旱型花生的主要根系分布特征; 土壤水分亏缺条件下, 花生主要通过增加深层土壤内根长、根系表面积和体积等形态特性调节植株对水分的利用。     

Assessment of earthworm burrowing efficiency in compacted soil with a combination of morphological and soil physical measurements

Summary Column experiments were carried out to quantify the effect of earthworms on compacted soil. The earthworms (Lumbricus terrestris) were able to burrow into soil which was artificially compacted to a pore volume as low as 40%; they may also penetrate an artificial plough pan deep in the soil. The effect of the burrowing activity of Lumbricus terrestris was quantified by measuring hydraulic conductivities and infiltration rates through the whole soil column (19 cm wide, 40 cm long). Morphological parameters, mainly the vertically projected burrow depth, were correlated with the saturated hydraulic conductivity. The amount of casts deposited by Lumbricus terrestris on the soil surface increased with the degree of soil compaction. The bulk density of casts was always less than that of the original soil.