Yvan Capowiez

Local Adaptation and Gene-For-Gene Coevolution in a Metapopulation Model

In several reciprocal cross-infection experiments parasites were found to be significantly more adapted to their local host populations than to hosts from distant populations. We developed a metapopulation model, taking explicit account of both population densities and gene frequencies, to determine the influence of ecological and genetical parameters on the local adaptation of the parasites and on the spatial distribution of resistance and virulence genes. Our results point to the predominant effect of ecological parameters such as parasite growth rate and host and parasite migration rates on coevolutionary outcomes. In particular, the parasites are more likely to be adapted to their local host population than to allopatric hosts when the parasite migration rate is larger than the host migration rate. The opposite should be observed whenever hosts migrate more than parasites.

3D reconstruction and quantification of macropores using X-ray computed tomography and image analysis

Axial X-ray Computed tomography (CT) scanning provides a convenient means of recording the three-dimensional form of soil structure. The technique has been used for nearly two decades, but initial development has concentrated on qualitative description of images. More recently, increasing effort has been put into quantifying the geometry and topology of macropores likely to contribute to preferential now in soils. Here we describe a novel technique for tracing connected macropores in the CT scans. After object extraction, three-dimensional mathematical morphological filters are applied to quantify the reconstructed structure. These filters consist of sequences of so-called erosions and/or dilations of a 32-face structuring element to describe object distances and volumes of influence. The tracing and quantification methodologies were tested on a set of undisturbed soil cores collected in a Swiss pre-alpine meadow, where a new earthworm species (Aporrectodea nocturna) was accidentally introduced. Given the reduced number of samples analysed in this study, the results presented only illustrate the potential of the method to reconstruct and quantify macropores. Our results suggest that the introduction of the new species induced very limited chance to the soil structured for example, no difference in total macropore length or mean diameter was observed. However. in the zone colonised by, the new species. individual macropores tended to have a longer average length. be more vertical and be further apart at some depth. Overall, the approach proved well suited to the analysis of the three-dimensional architecture of macropores. It provides a framework for the analysis of complex structures, which are less satisfactorily observed and described using 2D imaging

X-ray tomographic and hydraulic characterization of burrowing by three earthworm species in repacked soil cores

To test the assumption that different earthworm species make differently-structured burrow systems that can affect water permeability, the burrow systems made by adults of three earthworms species (Lumbricus terrestris, Nicodrilus giardi and Octalasion lacteum) were studied in repacked soil cores (20 cm diameter by 40 cm high) under controlled conditions of temperature and humidity. Five core were used for each species and one adult earthworm was incubated per soil core over 21 days. Three-dimensional reconstruction (skeletons and volumetric images) of the burrow systems was performed and characterized after computer analysis of the 2D-slices from X-rays tomography. Total burrow length, number of burrows, branching rate, connectivity, burrow continuity, mean verticality and mean sinuosity of the burrows were measured from the skeletons. Differences were found between species in burrow characteristics: only one vertical and continuous burrow with large diameter for L. terrestris; long and complex burrow systems with branching burrows for N. giardi and O. lacteum. Moreover, 3D volume reconstruction of burrow systems enabled determination of the total volume, the distributed burrow volume as a function of depth and, using the tools of mathematical morphology, the pore size distribution of burrows. Results showed that the burrow volume of N. giardi was the highest; the three species had a unimodal pore size distribution centered on 8, 7 and 5 mm in diameter for L. terrestris, N. giardi and O. lacteum, respectively. Saturated hydraulic permeability was measured for each core and predicted using a water flow model. Significant differences between species were found, the burrow systems made by N. giardi being the most efficient for water flow. Differences in hydraulic properties between species resulted mainly from the difference in burrow continuity, the presence of openings at the soil surface and the burrow depth penetration.

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

Abstract 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.

Pesticides and earthworms. A review

Earthworms provide key soil functions that favour many positive ecosystem services. These services are important for agroecosystem sustainability but can be degraded by intensive cultural practices such as use of pesticides. Many literature reports have investigated the effect of pesticides on earthworms. Here, we review those reports to assess the relevance of the indicators of earthworm response to pesticides, to assess their sensitivity to pesticides, and to highlight the remaining knowledge gaps. We focus on European earthworm species and products authorised in Europe, excluding natural compounds and metals. We consider different organisation levels: the infra-individual level (gene expression and physiology), the individual and population levels (life-history traits, population density and behaviour) and the community level: community biomass and density. Our analysis shows that earthworms are impacted by pesticides at all organisation levels. For example, pesticides disrupt enzymatic activities, increase individual mortality, decrease fecundity and growth, change individual behaviour such as feeding rate and decrease the overall community biomass and density. Insecticides and fungicides are the most toxic pesticides impacting survival and reproduction, respectively.

B-type esterases in the snail Xeropicta derbentina : An enzymological analysis to evaluate their use as biomarkers of pesticide exposure

The study was prompted to characterize the B-type esterase activities in the terrestrial snail Xeropicta derbentina and to evaluate its sensitivity to organophosphorus and carbamate pesticides. Specific cholinesterase and carboxylesterase activities were mainly obtained with acetylthiocholine (K(m)=77.2 mM; V(max)=38.2 mU/mg protein) and 1-naphthyl acetate (K(m)=222 mM, V(max)=1095 mU/mg protein) substrates, respectively. Acetylcholinesterase activity was concentration-dependently inhibited by chlorpyrifos-oxon, dichlorvos, carbaryl and carbofuran (IC50=1.35x10(-5)-3.80x10(-8) M). The organophosphate-inhibited acetylcholinesterase activity was reactivated in the presence of pyridine-2-aldoxime methochloride. Carboxylesterase activity was inhibited by organophosphorus insecticides (IC50=1.20x10(-5)-2.98x10(-8) M) but not by carbamates. B-esterase-specific differences in the inhibition by organophosphates and carbamates are discussed with respect to the buffering capacity of the carboxylesterase to reduce pesticide toxicity. These results suggest that B-type esterases in X. derbentina are suitable biomarkers of pesticide exposure and that this snail could be used as sentinel species in field monitoring of Mediterranean climate regions.

Earthworm behaviour as a biomarker – a case study using imidacloprid

To determine whether earthworm behaviour can be used as a biomarker for pollutant effects, an experiment was set up using ®Confidor (imidacloprid) as a model pesticide. The burrowing behaviour of two earthworm species ofdifferent ecological types (the anecic Aporrectodea nocturna and the endogeic Allolobophora icterica) was studied using 2D terraria. The activity of two classical biochemical markers (AchE and GST activities) was also measured. Imidacloprid had no effect on the biochemical markers in either earthworm species whatever the concentration (0.01, 0.1 and 1 ppm). In contrast, earthworm behaviour changed dramatically in the presence of imidacloprid. Following one week of exposure to the pesticide at two different concentrations (0.5 and 1 ppm), burrow length, the rate of burrow reuse and the distance covered decreased for both species. We conclude that earthworm behaviour is a sensitive biomarker that can be studied with cheap and simple apparatus such as 2D terraria.

X-ray computed tomography to quantify tree rooting spatial distributions

Poor root development due to constraining soil conditions could be an important factor influencing health of urban trees. Therefore, there is a need for efficient techniques to analyze the spatial distribution of tree roots. An analytical procedure for describing tree rooting patterns from X-ray computed tomography (CT) data is described and illustrated. Large irregularly shaped specimens of undisturbed sandy soil were sampled from Various positions around the base of trees using field impregnation with epoxy resin, to stabilize the cohesionless soil. Cores approximately 200 mm in diameter by 500 mm in height were extracted from these specimens. These large core samples were scanned with a medical X-ray CT device, and contiguous images of soil slices (2 mm thick) were thus produced. X-ray CT images are regarded as regularly-spaced sections through the soil although they are not actual 2D sections but matrices of voxels similar to 0.5 mm x 0.5 mm x 2 mm. The images were used to generate the equivalent of horizontal root contact maps from which three-dimensional objects, assumed to be roots, were reconstructed. The resulting connected objects were used to derive indices of the spatial organization of roots, namely: root length distribution, root length density, root growth angle distribution, root spatial distribution, and branching intensity. The successive steps of the method, from sampling to generation of indices of tree root organization, are illustrated through a case study examining rooting patterns of valuable urban trees

Differences in burrowing behaviour and spatial interaction between the two earthworm species Aporrectodea nocturna and Allolobophora chlorotica

Abstract To study intraspecific and interspecific interactions between different ecological types of earthworm, the burrowing behaviour of two earthworm species (the anecic earthworm Aporrectodea nocturna and the endogeic earthworm Allolobophora chlorotica) was observed in a microcosm. Earthworms were either alone in the microcosm, together with a conspecific earthworm, or with an earthworm of the other species. Observations under red light, including those of the position of the animals and the burrow dug, were recorded 4 times a day for 8 days and provided the data needed to reconstruct the probable trajectories of each earthworm. Differences in movement and burrowing behaviour were observed. Comparisons between the two species confirmed the expected behaviour of each ecological type: A. nocturna reused its burrow system regularly, whereas A. chlorotica rarely did. Moreover, it was shown that A. chlorotica burrowed less and explored a smaller surface when in the presence of A. nocturna. Besides, A. nocturna burrowed less and explored a smaller surface when in the presence of another A. nocturna. If these interactions occur under natural conditions, they could affect the structure of the burrow systems of the earthworm species examined.

Burrow systems made by Aporrectodea nocturna and Allolobophora chlorotica in artificial cores: morphological differences and effects of interspecific interactions

Burrow systems of earthworms contained in artificial cores were analysed through X-ray computed tomography and 3D skeleton reconstructions. Gas diffusion experiments were carried out on these cores to characterize soil transfer properties associated with the different burrow systems. Three types of cores were studied: cores in which Aporrectodea nocturna, an anecic earthworm was introduced (treatment 1), cores in which Allolobophora chlorotica, an endogeic earthworm was introduced (treatment 2) and cores that contained both species (treatment 3). Comparisons of the characteristics of the burrow systems of treatment 1 and 2 show important differences: the burrow system of A. nocturna comprises fewer burrows, which are longer, less branched, more vertical and have a lower sinuosity. The burrow system of A. chlorotica is characterized by lower continuity, which however did not result in a lower soil diffusivity. To study the burrow systems made by the two species in the same core (treatment 3), a separation that was based on differences in pore diameter between the two species and that takes into account the burrow orientation was designed. This separation was proven to be efficient since it resulted in low percentages of errors (around 10%) when applied to the burrow systems of treatments 1 and 2. Comparison of the burrow systems from treatments 1 and 3 demonstrated that the burrow system of A. nocturna was influenced by the presence of A. chlorotica: in treatment 3, A. nocturna made burrows that were smaller, more vertical and less branched. However, these interactions have to be confirmed under natural conditions.