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International audience; Root-knot nematodes are a growing concern for vegetable producers, because chemical nematicides are gradually disappearing. Alternative techniques based on agronomic practices are needed to solve the problem. This review analyzes the most recent studies related to these techniques and their combinations and identifies the most effective ones. Based on an agronomic point of view, the analysis focuses on a description of agricultural factors and practices, rather than on biological processes. Several alternative techniques are considered, including sanitation, soil management, organic amendments, fertilization, biological control and heat-based methods. We analyzed the effects of each practice and interactions among techniques and found large variations among studies. Many practices are only partially effective for nematode control; thus, combining control methods in a systemic analysis presents a challenge. We outline such an ongoing systemic approach and identify key future research studies.

Nematodes belonging to the genus Meloidogyne are the most ubiquitous and widespread plant-parasitic nematodes. They occur worldwide, are polyphagous and can parasitize most cultivated plants leading to reduced crop yields. They are especially harmful in developing countries because of the lack of suitable and feasible management strategies. Among all the control practices (chemicals, physical techniques, cultural practices, resistance), the use of natural enemies as biological control agents is the most recently developed. Pasteuria penetrans which is an obligate Gram-positive, endospore-forming bacterium, is perhaps the most promising plant-parasitic nematode biocontrol agent. Despite much research conducted on prey–predator interactions (host–parasite specificity, mechanisms of attachment, field efficacy), the influence of the soil environment on host–parasite interactions is poorly understood even when the soil appears to be the key factor. Beyond common studies on the influence of climatic conditions on the attachment of endospores of P. penetrans to nematodes, more knowledge about the systemic interactions between plants, soil water dynamics, soil texture and structure, and other biota on the parasitism of nematodes by P. penetrans would improve their utilization as biological control agents. The aim of this review is to analyze the literature dealing with the influence of the soil on nematode – P. penetrans interactions in order to suggest a helpful conceptual model based on partitioning the Pasteuria population in sub-populations according to their soil habitat (dispersible and non-dispersible aggregates, microporosity, macroporosity), not all of them being available for attachment and infection on nematodes. Such concerns should be taken into account by epidemiologists for improving biological management strategies based on the use of this bacterium.

The epidemiology of Pasteuria penetrans, parasite of Meloidogyne javanica, was studied in a field cultivated with an African egg-plant (Solanum aethiopicum) in Senegal. Abiotic and biological parameters were analyzed from 100 soil samples. Soil physical characteristics were studied with principal component analysis (PCA). Similarity between the parameters according to biotic and abiotic variables was established by hierarchical classification and corroborated by Discrimin analysis (AFD). The coarser fractions (> 200 mu m) were positively correlated to intensive irrigation, whereas finest fractions (< 20 mu m) and spores of P. penetrans were abundant in the areas moderately irrigated. Furthermore, the proportion of Pasteuria-infected juveniles was positively correlated to the presence of finest texture and high water holding capacity. The abundance of M. javanica was associated with coarse sands and potassium. Based on these results, laboratory experiments were attempted on the influence of irrigation and soil texture on the distribution of P. penetrans in cultivated and bare soils. The results showed that 53% of the spores inoculated were leached by water flow in the sandy soil, but only 14% in the sandy-clay soil and 0.1% in the clay soil. No nematodes survived in the clay soil, while the population increased both in the sandy and the sandy-clay soils. More M. javanica were infected by P. penetrans in the sandy-clay soil than in the sandy soil. Comparing different combinations of bare soils containing 1.1-57% clay, the best spore percolation/retention balance occurred in soils textured with 10-30% clay.

International audience; Soil is a component of primary importance in crop production, even if it is often neglected, or only regarded as a physical support for the growth of plants. However, with the increasing societal concerns for the sustainability of agriculture, soil must be considered as a living system. Its quality results from the multiple interactions among physicochemical and biological components, notably the microbial communities, primordial for soil function. Crops are threatened by soil-borne diseases. These are often difficult to control, because of the “hidden” status of the pathogens and also because of the absence, noxiousness or lack of efficacy of chemical treatments. In this context, there is a renewed interest for cultural practices such as the use of organic amendments. These practices have a strong influence on soil health, which takes into account plant health, through both changes of physicochemical characteristics and influence on soil microbial communities. Cultural practices are used to improve soil health, and can, in some cases, increase soil disease suppression. The effects of these practices on soil properties have been studied, but the relationship between these properties and soil suppressiveness has not been always very clear. Many different soil descriptors, either abiotic or biotic, have been used to describe the soil health and suppressiveness, but there is a lack of identified, reliable and consistent indicators. One aim of this review is to show that, despite the age of the soil health concept and all the studies that have been conducted, there are still no guidelines for assessment of soil quality from the plant health point of view. Obviously, the extreme diversity of situations makes any generalisation from a given case-study difficult. However, based on what has already been done in related fields, a methodology to search for indicators of soil health can be proposed. In this review we will present (i) how a healthy soil can be defined and what are the concepts hidden behind the words “soil health,” (ii) which cultural practices have been used to control soil-borne diseases and their limitations, (iii) which soil parameters have been measured when studying soil suppressiveness, and which relationships have been found between these parameters, and finally (iv) how these descriptors could become indicators of soil health, using appropriate analytical and statistical methods.

Summary In Senegal, a study was carried out in a field cultivated with African egg-plant ( Solanum aethiopicum ) in order (1) to determine soil size fractions infested both with Meloidogyne javanica and Pasteuria , (2) to compare the efficacy of soil fraction dispersion methods in the recovery of both organisms and (3) to estimate the concomitant distribution of the two organisms in the different soil fraction sizes. Samples were taken from four areas: severely-infested (M+) with M. javanica with a high percentage (I+) of Pasteuria -infected juveniles, severely-infested (M+) with a low percentage of infected juveniles (I−), minimally-infested (M−) with a high percentage of infected juveniles (I+) and minimally-infested (M−) with a low percentage of infected juveniles (I−). The samples were dispersed and fractionated using two different techniques in order to evaluate soil aggregation (size fractions >200, 50–200, 20–50 and 0–20 μm). The numbers of free entities of Pasteuria penetrans were estimated per size fractions. The proportion of Pasteuria -infected juveniles (biological detection of P. penetrans ) was estimated in order to correlate the infection rate to the distribution of free entities of P. penetrans . Regardless of sampling area, the 50–200 μm size fraction predominated, followed by largest particle (>200 μm) size. The distribution of Pasteuria spores in the soil was not linearly correlated to the distribution of Pasteuria -infected juveniles. The influence of the soil characteristics on the relationship between the two organisms is discussed.

Pasteuria penetrans, a bacterial parasite of plant-parasitic nematodes, is used to control root-knot nematode Meloidogyne spp. populations in vegetable crops. But its efficiency is variable, mostly because of the patchy distribution of the bacteria in arable fields. As the infective P. penetrans are non-motile bacteria in soil, abiotic soil factors can affect the bacteria–nematode relationships. An epidemiological study, conducted in a vegetable field, showed that abiotic factors such as irrigation, soil water holding capacity and texture, affected the efficiency of P. penetrans. A correspondence analysis between these abiotic factors and the density of P. penetrans spores in the soil, and the proportion of Meloidogyne javanica juveniles infected by the bacteria, revealed that irrigation affected directly the distribution of the spores in soil pores related to their passive transport by water flow. Laboratory experiments conducted on the passive transport of spores confirmed that intensive irrigation leached the spores down the soil profile and decreased the percentage of infected Meloidogyne juveniles.

The transport of the spores of Pasteuria penetrans was studied in three contrasted textured soils (a sandy, a sandy-clay and a clay soils), cultivated with tomato, inoculated with juveniles of Meloidogyne javanica and watered with 25 or 150 mm day−1. One month after inoculation of the nematodes, 53% of the spores inoculated were leached by water flow in the sandy soil but only 14% in the sandy-clay soil and 0.1% in the clay soil. No nematodes survived in the clay soil, while the population was multiplied both in the sandy and in the sandy-clay soils. But juveniles of M. javanica were more infected by P. penetrans in the sandy-clay soil than in the sandy soil. Comparing different combinations of bare soils containing 1.1–57% of clay showed that the best spore percolation and retention balance occurred in soils amended with 10–30% clay. However, the spore recoveries decreased when the soil was enriched with more than 30% clay. The role of clay particles on the extractability of spores and on their availability to attach to the nematode cuticle in the soil is discussed.

Differences in rates of infestation of root-knot nematode (Meloidogyne spp.) populations by the endospore-forming bacterium Pasteuria penetrans were not entirely due to inter/intra-specific variability of the organisms. Soil conditions, especially texture and chemical characteristics, are also involved. Focusing studies on vegetable crops, interactions between the occurrence of Meloidogyne spp. juveniles encumbered with spores of P. penetrans and some physico-chemical soil characteristics were analysed both between regions (West Africa, e.g. Burkina Faso and Senegal; South America, e.g. Ecuador; the Caribbean, e.g. Trinidad and Tobago; and Mediterranean Europe, e.g. Crete) and within Ecuador, Burkina Faso and Senegal. In Ecuador (clay or silty-clay soils), the mean proportion of infested juveniles was high (41.9%). In the sandiest soils, as in Senegal, there were very few infested juveniles (4.7%). In Crete, Burkina Faso, and Trinidad and Tobago, where the soils are siltier, the mean proportions of infested juveniles range between 14 and 24.4%. Multivariate analysis performed on the data from Senegal and Burkina Faso revealed that a significant increase of the mean clay content (1.4 and 10.5%, respectively) improved the mean proportions of infested juveniles (from 1.6 to 42.2% and from 10.7 to 79.4%, respectively). Influences of the soil texture and structure on the availability of the spores of P. penetrans to infest the juveniles of Meloidogyne spp. are discussed.

The conclusions of a collaborative study of the occurrence and importance of root-knot nematodes (RKN, Meloidogyne spp.) and of their control agent, Pasteuria penetrans, in parts of Europe, Africa, South America and the Caribbean are presented. Rootknot nematodes were estimated to reduce the yields of a wide range of horticultural crops by > 25% in Ecuador, Malawi and Tanzania, and by ca 10% in Trinidad and Tobago. The greatest proportion of infected crops were observed in Ecuador (205 of 207) and the least in Trinidad and Tobago (70 of 174). The mean gall index was greatest in Ecuador (5.5). Levels of galling were least in Senegal (1.6), even though 89% of crops were infested and virulent M. mayaguensis was widespread. In all countries, M. incognita and M. javanica were the most abundant species, but M. hispanica occurred widely in Burkina Faso, even in newly cultivated areas in the Sahile. Several new esterase phenotypes were detected, especially in Ecuador and Malawi. Juveniles (J2) collected from the soil during the surveys were examined for attached spores of P.penetrans. It was widespread (20 to 60% of RKN populations), except in Malawi and Tanzania (< 10% were infected), and was found for the first time in Crete (Greece). Generally, < 50% of the J2 carried spores. The occurrence of P.penetrans was sometimes correlated with soil type e.g., in Senegal it was least frequent in sandy soils. Laboratory assays of the binding of spores of isolates of P.penetrans to populations of RKN indicated large variations in specificity and substantial interactions; differences between populations within a species of RKN were sometimes almost as great as those between species. In microplot trials in which an "exotic" isolate of P.penetrans was introduced (ca 103 spores per g soil), its incidence was not increased by increasing the frequency or intensity of the growing of RKN-susceptible crops. However, in two such trials at sites in Tanzania and Ecuador naturally infected with P.penetrans, there were large increases in the proportions of spore-encumbered J2 (up to 100% encumbered) and in the yields of spores (up to 3.3 × 106 spores per mg dry root) in those plots amended with an "exotic" isolate. In these plots, numbers of J2 in the soil were decreased and damage by RKN was suppressed; gall indices were decreased (from > 8 to < 3) and yields were increased (by up to 30%). No such changes were observed in the unamended control plots. Increased suppression of RKN was also observed in a field trial, even in plots where RKN-susceptible and non-host crops were alternated. Increased suppression following amendment with the "exotic" isolate of P.penetrans was not observed at sites not previously infected with P.penetrans. Regression analysis of the results from the microplot and field trials indicated that tomato yields were decreased by > 5% for every increase of one in the gall index. Yields were increased by alternating tomato with leguminous crops in some trials, but not in others. It is proposed that, in natural infections, mutual selection produces a dynamic balance between the P.penetrans and the RKN whereby levels of infection are rarely suppressive. However, the introduction of an "exotic" isolate of P.penetrans, with a different attachment profile, can disturb this balance, resulting in a greatly increased proportion of infected J2 and females, increased yields of spores and more suppression of RKN populations. Importance des nématodes à galles tropicaux (Meloidogyne spp.) et facteurs affectant l'utilité de Pasteuria penetrans, agent de contrôle biologique - Ce travail présente les conclusions d'une étude, menée en collaboration par plusieurs équipes de recherche, sur la présence et l'importance des nématodes phytoparasites du genre Meloidogyne et de leur parasite bactérien, Pasteuria penetrans, dans certains pays d'Europe, d'Afrique, d'Amérique du Sud et des Caraïbes. Les réductions de rendement de cultures maraîchères très diverses dues à ces nématodes atteignent 25% en Equateur, Malawi et Tanzanie, et 10% à Trinidad et Tobago. La plus forte proportion de parcelles infestées a été rencontrée en Equateur (205 sur 207) et la plus faible à Trinidad et Tobago (70 sur 174). C'est en Equateur que l'indice de galles moyen (égal à 5,5) était le plus élevé. Même si cet indice était faible en moyenne au Sénégal (1,6), 89% des cultures étaient infestées dans ce pays, en grande partie par l'espèce virulente M. mayaguensis. M. incognita et M. javanica sont les plus répandues dans tous les pays prospectés. Cependant, M. hispanica est très répandu au Burkina Faso, même dans des zones récemment cultivées en maraîchage en région sahélienne. Plusieurs phénotypes estérasiques nouveaux ont été détectés, spécialement en Equateur et au Malawi. Les juvéniles de second stade (J2) extraits des échantillons de sol collectés lors des prospections ont été examinés pour détecter la présence de spores de P.penetrans sur leur cuticule. Trouvé pour la première fois en Crète (Grèce), P.penetrans est très répandu dans les autres pays prospectés, infestant 20 à 60% des populations de Meloidogyne spp., sauf au Malawi et en Tanzanie où moins de 10% des populations sont atteintes. Le plus souvent, moins de 50% de J2 portent des spores bactériennes. Le taux de parasitisme des J2 par P. penetrans est influencé par les types de sols, comme par exemple au Sénégal où il est très faible dans les sols sableux grossiers. Des expériences en laboratoire portant sur l'attachement de spores de divers isolats de P. penetrans à des populations de Meloidogyne spp. ont révélé une grande variation de la spécificité et des interactions; les différences observées pour diverses populations d'une même espèce de Meloidogyne sont parfois presque aussi marquées que celles observées entre espèces. L'incidence parasitaire d'un isolat "exotique" de P. penetrans introduit dans des microparcelles (ca. 103 sporespar g. de sol) n'a pas été accrue par l'augmentation de la fréquence ou de la densité de plantation des cultures sensibles à Meloidogyne spp. employées. Cependant, dans deux microparcelles natullement infestées en P.penetrans, situées en Equateur et en Tanzanie, l'introduction d'un isolat "exotique" s'est traduite par un accroissement de la proportion de J2 infestés (jusqu'à 100%) et de la production de spores ((jusqu'à 3, 3 × 106 spores par mg [poids sec] de racines), d'une diminution de la population de J2 dans le sol, et d'une disparition des dégâts racinaires; les indices de galles moyens ont diminué (de plus de 8 à moins de 3) et les rendements des cultures ont augmenté (jusqu'à 30%). De tels changements n'ont pas été observés dans des sols non amendés en isolats "exotiques" de P.penetrans. Un meilleur contrôle des populations de Meloidogyne spp. a également été observé au champ, même lorsque la séquence culturale alternait des cultures sensibles et des cultures non-hôtes du nématode. Dans des parcelles non infectées en P.penetrans, la baisse des populations de Meloidogyne spp. n'a pas été observée après introduction d'un isolat "exotique" de la bactérie. Des analyses de régression portant sur les données obtenues en microparcelles ou au champ montrent que les rendements en tomate diminuent de plus de 5% chaque fois que l'indice de galle augmente d'une unité. Ces rendements ont parfois été améliorés lorsque des cultures de légumineuses alternaient les cultures de tomate. Ainsi, sur la base des analyses nématologiques et agronomiques faites en fin de cycles culturaux, il est suggéré que, dans les cas d'infestations naturelles en P.penetrans, des sélections mutuelles entraîneraient un équilibre dynamique entre les populations de la bactérie et celles du nématode, représentatif d'une densité-dépendance retardée. En revanche, l'introduction inondative d'isolats "exotiques" de P.penetrans, aux propriétés parasitaires différentes de celles des populations natives, pourraient rompre temporairement cet équilibre en faveur d'un accroissement de la proportion de nématodes (J2 et femelles) infestés et de la production de spores bactériennes, ainsi qu'un meilleur contrôle des populations de Meloidogyne spp. La capacité des populations de P.penetrans à survivre dans les sols et à contrôler durablement les populations de Meloidogyne spp. dépendraient de la spécificité entre les organismes, des propriétés des sols et des systèmes de culture.

Two strains (ORS 18692 S7 and ORS 18692 S5) of the nematophagous fungus Arthrobotrys oligospora have been isolated for the first time in Senegal. In vitro, both strains trapped 98% of groups of 7-day-old Meloidogyne mayaguensis juveniles within 48 h. Their optimal growth occurred at 25-30 C and at a pH of 5.6, but salinity inhibited their development. In order to test the fungi on M. mayaguensis in pots and in mixed populations of Meloidogyne spp. on tomato in the field, the fungus was incorporated into compost blocks before sowing and subsequent transplanting. In pot experiments during the cool season (28 C), both strains reduced nematode populations and stimulated seedling growth. In field trials during the hot season (35 C), both strains were efficient in reducing Meloidogyne populations. The feasibility of combining the use of compost blocks with the introduction of A. oligospora is discussed. Pour la premie re fois, deux souches du champignon ne matophage Arthrobotrys oligospora (souches ORS 18692 S5...

The susceptibility of two acacias (Acacia holosericea and A. seyal) against the root-knot nematode Meloiodogyne javanica has been studied. The nematode was inoculated at different rates (1000, 5000 and 10,000 second stage juveniles per seedling). In both acacias, M. javanica inhibited seedling height, shoot and root biomass and the density of root tips per mg of root. In terms of nematode development, the increase of M. javanica was greater on A. holosericea than on A. seyal when the inoculum rate was 1000 J2 per seedling indicating that A. holosericea is a better host for M. javanica than A. seyal.