Dans les paysages agricoles, les habitats non cultivés constituent une source importante pour l’entomofaune auxiliaire de cultures. Il est donc important d’adapter la gestion de ces habitats afin de bénéficier au mieux des services fournis par la diversité de ces auxiliaires. L’objectif de cette thèse est de comprendre comment la gestion des bandes enherbées permet d’améliorer le service du contrôle biologique délivré par les carabes. Nous avons d’abord examiné par quels mécanismes, les changements dans la diversité fonctionnelle des carabes augmentent leur capacité de suppression de ravageurs. Ainsi, a été comparée au laboratoire, l’efficience de prédation sur une population de proie, par des assemblages de carabes qui ont différents niveaux de diversité de taille. Par ailleurs, à travers une expérimentation mise en place dans des champs de céréales, nous avons étudié les effets de cinq modes de gestion des bandes enherbées sur la structure végétale de ces habitats non cultivés, et par conséquent sur la structure des communautés carabiques. Nos résultats révèlent l’absence d’une partition de ressources entre les carabes de tailles différentes. La quantité et de la taille de proies consommées augmente avec la taille des carabes. La suppression des proies a ainsi été fortement augmentée par la présence des carabes de grande taille dans les assemblages. Ces résultats suggèrent que l’augmentation de la diversité de taille chez les carabes augmente leur efficience de prédation par l’effet d’échantillonnage. Nous avons mis en évidence une redondance fonctionnelle entre les espèces de carabes. Cette redondance permet de réduire la variabilité spatio-temporelle dans la fonction de régulation exercée sur les populations de ravageurs. L’importance des bandes enherbées en tant qu’habitat pour plusieurs espèces de carabes importantes pour le contrôle biologique, a été mise en évidence. La gestion influence les carabes seulement dans les bandes enherbées. La diversité fonctionnelle des plantes au sein de ces bandes enherbées a été identifiée comme le principal facteur favorisant la richesse spécifique des carabes. L’activité-densité des carabes a, par ailleurs, été favorisée par une végétation peu hétérogène et un pourcentage élevé de sol nu. Les carabes de tailles différentes divergent dans leur réponse à la structure de la végétation. Cette divergence semble être liée aussi à la variation d'autres traits d'histoire de vie des espèces (i.e. régime alimentaire). Les connaissances et les résultats produits dans le cadre de ce travail de thèse, serviront au développement d’indicateurs mesurant l’impact des pratiques de gestion sur le contrôle biologique exercé par les carabes / In the agricultural landscapes, non cropped habitats are an important source for the beneficial entomofauna of crops. It is therefore important to adapt the management of these habitats to best benefit of the services provided by the diversity of this beneficial entomfauna. The objective of this thesis is to understand how field margin management could improve the biological control service delivered by ground beetles. First, we examined by which mechanisms, changes in the functional diversity of ground beetles increases their capacity of pest suppression. Thus, efficiency of predation by ground beetle assemblages, which have different levels of body size diversity, on a prey population was compared in the laboratory. Also, through an experiment installed in cereal fields, we studied the effects of five field margin management on the vegetation structure of these non cropped habitats, and therefore on the structure of ground beetle communities. Our results revealed the absence of resource partitioning among the different-sized ground beetles. The amount and size of prey consumed increased with increasing ground beetle body size. Prey suppression was, therefore, strongly strengthened by the presence of large ground beetles in the assemblages. These results suggest that increasing ground beetles size diversity increases their predation efficiency through the sampling effect. We have showed a functional redundancy among species of ground beetles. This redundancy could reduce the spatial and temporal variability in pest suppression by ground beetles. Our study showed the importance of the field margins as a habitat for many ground beetle species that are important for biological control. Management influenced ground beetles only in the field margin. Plants functional diversity within these field margins has been identified as the main factor promoting the species richness of ground beetles. However, activity-density of ground beetles was promoted by less heterogeneous vegetation and high percentage of bare ground. The responses to the vegetation structure vary between ground beetles with different body sizes. This diversity in ground beetles responses to the vegetation structure appears to be also related to variation in other life history traits (i.e. diet). The knowledge and results produced as part of this thesis will be used in the development of indicators measuring the impact of management on biological control delivered by ground beetles

Augmenter la diversité végétale au sein même du champ permet de réguler les populations de ravageurs dans de nombreux agroécosystèmes. Les mélanges variétaux (diversité intraspécifique) ou les associations de cultures avec une plante compagne (diversité interspécifique) sont considérées comme des pratiques agroécologiques prometteuses pour les systèmes de culture à bas intrants ou l'agriculture biologique. En effet, ces pratiques favorisent de nombreux services écosystémiques tels que la régulation des ravageurs, des maladies ou des adventices, ainsi que la fertilisation azotée. Cependant, le potentiel de régulation des ravageurs du blé par la combinaison de ces deux pratiques de diversification n'a pas encore été étudié.Nous avons combiné ces deux pratiques dans le cadre d'expérimentations menées en plein champ et sur deux saisons de culture, afin d'examiner leurs impacts sur les populations de pucerons et d'ennemis naturels. Nous avons également évalué le potentiel de régulation des ravageurs en mesurant les taux de prédation de proies sentinelles.La combinaison des diversités intra- et interspécifique n'est pas plus performante pour réduire les populations de pucerons que les pratiques prises séparément. L'association de culture blé-trèfle tend à être moins infestée par les pucerons, tandis que le mélange variétal est plus infesté que la variété la moins sensible. Les variations annuelles des conditions climatiques impactent fortement le développement du blé et du trèfle, ainsi que la date d'apparition du pic de puceron. Le rendement du blé, ainsi que le taux d'azote du grain sont réduits par l'association de culture par 7 à 10%, mais pas par le mélange variétal. La présence d'un couvert de trèfle dans les champs de blé, semble avoir favorisé la biodiversité fonctionnelle, particulièrement les ennemis naturels tels que les carabes, mais pas le mélange variétal. Les résultats sont variables selon la famille d'arthropodes concernée et leur position au sein du couvert végétal (au sol ou dans le feuillage). Le couvert de trèfle et le champ ont influencé la composition de la communauté de carabes prédateurs. Les taux de prédation des proies sentinelles n'ont pas été impactés par les pratiques de diversifications.En laboratoire, nous avons évalué comment l'association du blé avec des légumineuses (trèfle ou pois) pouvait modifier le comportement du puceron du blé Sitobion avenae en terme de location de sa plante hôte et du développement de la population. Les pucerons ont résidé moins de temps sur le blé quand il était associé à du trèfle. Les populations de pucerons se sont moins développées dans les associations du blé avec une légumineuse par rapport à du blé seul, mais si l'on prend en compte la biomasse du blé, seulement l'association blé-trèfle a considérablement réduit les densités de pucerons sur le blé. Ainsi l'espèce associée et sa densité sont des paramètres importants qui devraient être pris en compte dans les études sur la diversité interspécifique, car ils pourraient expliquer la grande variation dans les résultats rapportés par les analyses bibliographiques.Nos résultats suggèrent qu'augmenter la diversité cultivée au sein du champ peut aider à réguler les pucerons dans une certaine mesure, mais la combinaison des deux pratiques de diversification ne résultent pas en un trade-off entre la régulation des ravageurs et les performances agronomiques particulièrement attractifs pour les agriculteurs. / Increasing intrafield plant diversity has been shown to regulate pest populations in various agroecosystems. Polyvarietal mixtures of a crop species (intraspecific diversity) or associations of a crop and a companion plant (interspecific diversity) are both considered as promising agroecological practices for low-input or organic agriculture systems by providing several ecosystem services such as pest, disease and weed control, and nitrogen fertilization. However, combining both diversification practices has not been studied yet in perspective of winter wheat pest control.In organic field experiments over two growing seasons, we combined both practices and examined the direct impact on aphid and natural enemy populations and on wheat production. We also investigated the potential pest regulation service through the assessment of the rate of predation by using sentinel preys.Results show that combining intra- and interspecific diversity did not outperform each practice individually in reducing aphid populations, thus not clearly showing synergetic effects. Taken separately, intercropping tended to have lower aphid infestation, while it the cultivar mixtures was more infested by aphids than the least susceptible cultivar. Yearly variation in climatic conditions strongly impacted wheat and clover development, as well as the appearance of aphid peaks. Wheat yields and grain nitrogen content were reduced in intercropping by 7 to 10%, but not in cultivar mixtures. Functional biodiversity, especially natural enemies such as ground beetles, tended to be positively correlated to the presence of a clover cover in the wheat fields (interspecific diversification), but did not respond to the wheat cultivar mixture (intraspecific diversification). Results varied according to the family of arthropods concerned and their position within the vegetation layer (ground dwelling or foliage dwelling arthropods). The cover of white clover and the field context influenced the community composition of predatory ground dwelling beetles. Rates of predation on sentinel preys were not influenced by any of the diversification practices.Under laboratory conditions, we evaluated how combining wheat and legumes (clover or pea) modifies the behaviour of the cereal aphid Sitobion avenae in terms of host-plant location, and population growth. We observed that aphids’ residence time on wheat was decreased when this host-plant was intercropped with clover. At the population level, wheat-legume intercrops reduced the number of aphids on wheat plants compared to wheat sole crops but if we take into account plant biomass, only intercropping clover with wheat significantly reduced aphid densities on wheat. The species used as non-host plants and their density are important parameters that should be taken into account in studies on intercropping systems and that may explain the large variability in the results observed in the literature.Our findings suggest that intrafield diversification may regulate wheat aphids to some extent, but combining the two diversification practices did not result in an interesting trade-off between pest regulation and wheat production in real farming conditions.

La lutte biologique par conservation des habitats est une voie pour réduire la dépendance et l’impact des traitements insecticides qui sont nombreux et toxiques. Elle est particulièrement appropriée pour les vergers d’agrumes qui comprennent, souvent, de grandes surfaces non cultivées. Les auxiliaires présents en plus grand nombre dans un enherbement sont alors plus facilement mobilisables lors d’attaques massives de ravageurs sur les arbres. Ce travail de thèse a pour objectif d’évaluer l’impact des modes de gestion de l’enherbement (fauche, broyage, travail du sol et herbicide) sur les communautés végétales et les populations d’auxiliaires au sein du verger, en s’intéressant à deux communautés évoluant à deux échelles spatiales distinctes : les Phytoseiidae et les Coccinellidae. Nos résultats ont montré une différenciation fonctionnelle des communautés végétales soumises à différents modes de gestion dans le temps. Les Phytoseiidae sont présents en abondances comparables dans toutes ces communautés végétales mais sont cependant influencés par l’abondance dans le couvert des espèces végétales portant des densités de trichomes modérés et des domaties. De même, les abondances de Coccinellidae sont positivement corrélées à l’abondance des Poaceae sur la parcelle alors que les modalités de gestion de l’enherbement testées n’ont pas montré d’effet significatif. Ces connaissances ont pu être mobilisées et articulées pour l’élaboration d’un prototype d’outil d’aide à la décision contribuant à la conception de vergers agroécologiques. Il permet l’évaluation ex ante de scénarios de pratiques de gestion de l’enherbement en vue d’optimiser la lutte biologique par conservation au sein des vergers d’agrumes. Nous avons choisi une approche fonctionnelle qui permettra de généraliser facilement l’outil à d’autres cultures moyennant un élargissement de la base de données comportant les traits fonctionnels aux espèces végétales non renseignées et une bonne connaissance des affinités des auxiliaires que l’on cherche à favoriser / Biological control through habitat conservation is a way to reduce the reliance on and the impact of insecticide treatments that are numerous and toxic. It is particularly suitable for citrus orchards which include large areas often uncultivated. The natural enemies present in greater number in a vegetal cover are then more easily mobilized during massive attacks of pests on trees. The aim of this thesis is to evaluate the impact of different weed management methods (mowing, milling, tillage and herbicide) on plant communities and natural enemies populations within the orchard, focusing on two communities that evolve at two distinct spatial scales: the Phytoseiidae and the Coccinellidae. Our results showed a functional differentiation of vegetal communities managed with different weed management methods over time. Phytoseiidae are present in comparable abundances in all these plant communities but are influenced by the abundance of plant species with densities of moderate trichomes and domaties in the vegetal cover. Similarly, the abundances of Coccinellidae are positively correlated with the abundance of Poaceae species on the plot whereas the weed management modalities have no significant impact. This knowledge has been mobilized and articulated for the development of a decision-making tool prototype contributing to the design of agro-ecological orchards. It allows ex ante evaluation of scenarios of weed management practices in order to optimize biological control by conservation in citrus orchards. We have chosen a functional approach which will easily allow to generalize the tool to other crops by expanding the database with functional traits to non-informed plant species and a good knowledge of the affinities of natural enemies to promote

Le lin, principale plante à fibres cultivée en France possède un intérêt industriel pour la qualité de ses fibres. Les cultures de lin sont régulièrement attaquées par un champignon tellurique, Fusarium oxysporum f. sp. lini (Fol), responsable des plus grandes pertes dans les cultures de lin. Les PGPR (Plant Growth Promoting Rhizobacteria) sont des bactéries réputées pour leurs capacités à améliorer la croissance et le développement des plantes, mais également pour leur pouvoir compétiteur au sein de la rhizosphère et leur aptitude à induire une réponse immunitaire chez les plantes. Parallèlement, l’utilisation de SDP (Stimulateur des Défenses des Plantes), molécules capables d’éliciter les mécanismes de défenses des plantes est une autre alternative pour limiter l’utilisation des pesticides. Dans le cadre de ce projet mené sur deux variétés de lin, Aramis et Mélina, il a pu être montré que Fol induisait un remodelage de la paroi au niveau des racines et des tiges, impliquant les hémicelluloses et les pectines, seulement deux jours après inoculation avec le champignon. L’utilisation de la souche ATCC 6633 de Bacillus subtilis comme agent de biocontrôle a permis de réduire de façon significative l’apparition des symptômes de la fusariose. Il a également été montré qu’en plus d’avoir un effet fongicide sur Fol, cette bactérie est capable d’induire l’expression de deux gènes de défense (Pathogenesis-Related) codant pour une β-(1,3)-glucanase (PR-2) et codant pour une chitinase-like (CTL-10), de gènes impliqués dans la voie des phénylpropanoïdes (PHENYLALANINE AMONIA LYASES, PAL-3 et PAL-4) et dans le remodelage pariétal (PECTIN METHYLESTERASE-3, PME-3) au niveau racinaire. Des analyses biochimiques ont également permis de montrer que B. subtilis provoque des modifications se traduisant par un renforcement pariétal au niveau des tiges chez les deux variétés. Enfin, l’association de la PGPR avec une molécule élicitrice (pregnénolone sulfate) a eu un effet synergique sur l’expression de gènes de défense. / In France, flax (Linum usitatissumum) is a principal fibers crop. Fusarium oxysporum f sp lini (Fol), a soil-borne fungus, is responsible for the major losses in crop yield. PGPR (Plant Growth Promoting Rhizobacteria) are known for their abilities to promote plant growth and health. These bacteria are also good competitors in the rhizosphere and can induce a plant defense response. The use of compounds able to elicit plant defense mechanisms is also an alternative to limit the use of pesticides. In this project, it has been shown that F. oxysporum f. sp. lini induces only two days after inoculation cell wall remodeling in the root and the stem involving hemicelluloses and pectins on two flax varieties, Aramis and Mélina, . The use of the Bacillus subtilis strain ATCC 6633 as biocontrol agent significantly reduced fusarium wilt appearance. In addition to its antifungal effect against Fol, this bacteria is able to induce the expression of two Pathogenesis-Related genes coding for a β-(1,3)-glucanase (PR-2) and a chitinase-like (CTL-10), genes involved in the phenylpropanoid pathway (PHENYLALANINE AMONIA LYASES, PAL-3 and PAL-4) and also in cell wall remodeling (PECTIN METHYLESTERASE-3, PME-3) in the root. Biochemical analyses show that B. subtilis causes modifications resulting in cell wall reinforcement in the stem in both varieties. Finally, the association of B. subtilis with an elicitor (pregnenolone sulfate) had a synergistic effect on the expression of defense-related genes.

The use of biological control agents (BCAs) in the past has shown limited success as its application has often been done incorrectly, and in addition, management practices are rarely altered to incorporate BCAs. Criteria for the correct application of BCAs have been devised as part of the research, and companies selling these products may use the said criteria. Such application will ensure the correct BCAs are used and, more specifically, used under the correct conditions. The powdery mildew (PM) fungus is often seen to develop resistance to synthetic fungicides and, therefore, alternative control measures are required. BCAs as an alternative pose less risk to the environment, workers and the consumer. A pre-trial has been conducted with a range of BCAs to see if they can control powdery mildew (PM) in a greenhouse environment on hydroponically grown cucumber (Cucumis sativus L.) plants using the variety Baccara that has only a moderate tolerance to PM. The BCAs have been compared to the control (synthetic fungicide: Bravo). Comparative work includes Coyier's model, which has been modified and adapted for these trials to determine the percentage of leaf area covered by the PM infection. Furthermore, the number of fruit harvested per treatment, kilogram yield, total mass of yield and average fruit mass is also used to determine the efficacy of the BCAs as these factors have economic significance to commercial growers. The pre-trial showed promise until the fertigation computer failed, resulting in a nutrient shortage and imbalance, confirming that BCAs alone cannot control PM. Synthetic fungicides were applied until control of PM and plant nutrition was regained. BCAs were re-introduced and used until the end of crop production. The confirmation from the pre-trial has led to the inclusion of silicon in conjunction with the BCAs in the two subsequent trials (Trials 1 & 2). Silicon was applied with the BCAs as a foliar spray on a weekly basis. In trials 1 and 2, the cucumber variety, Palladium, with a high genetic tolerance to PM is used, as this variety is suited to form part of the holistic approach used for trials 1 and 2. Trial 1 showed that treatment A, containing Streptomyces griseovirdis and Streptomyces aureofaciens, had the highest yield. Both of these are bacterial BCAs and demonstrated their adaptability to varied climatic conditions, notably when low humidity was experienced. In treatment B, Trichoderma harzianum strains, Rifai and Uppington, show the slowest rate of PM development. In trials 1 and 2, the best actual PM control was obtained by two fungal based BCAs (Trial 1, treatment C was Ampelomyces quisqualis) and (Trial 2, treatment B was Trichoderma harzianum strains, Rifai and Uppington), showing that fungal BCAs have a place for this application, but the growth-enhancing properties of bacterial based BCAs make economic sense and would make them attractive to growers. Treatment A (Streptomyces spp.) had the most number of fruit for the entire growing period and the best overall yield (kg yield) again. Two of the BCA / silicon treatments have marginally better PM control compared to that of the control (E) treatment, although not statistically significant. Treatment E (control) has the highest average fruit mass in this instance but does not have the highest yield (kg yield) when compared to treatments A and B, possibly due to the growth-enhancing properties of most of these BCAs. Therefore, most of these BCA treatments give fairly inconsistent results that vary possibly according to season, humidity and temperature, making it difficult to predict their efficacy. Using combinations or weekly alternations of these BCAs with extremes of climatic adaptation will probably be the most reliable method of obtaining consistent results. Bacterial BCAs are shown to have lower humidity requirements and produce the most consistent results in terms of fruit number, yield and fruit mass and a combination of bacterial and fungal based BCAs would possibly be the best as this would control PM and yet still have the growth enhancing properties from the bacterial based BCAs. From the research, it can be said that some BCAs in trials 1 and 2 produce results similar to that of the control in terms of percentage leaf area covered by PM and some are shown to have improved yields. Results produced from certain BCA treatments are thus equal to the control; yet provide an environmentally friendly alternative to synthetic fungicides. Silicon is listed as a beneficial element rather than an essential element; however, literature claims it to be highly effective in treating PM in cucurbits. Results from trials 1 and 2 show that control of PM is possible in most cases, when a holistic approach is used. This approach includes a cucumber variety with a high PM tolerance, optimum nutrition, cultural practices and silicon in combination with the BCAs. A complete change of management practices is necessary to implement such a BCA program. / Agriculture, Animal Health & Human Ecology / M. Tech. (Nature Conservation)

The use of biological control agents (BCAs) in the past has shown limited success as its application has often been done incorrectly, and in addition, management practices are rarely altered to incorporate BCAs. Criteria for the correct application of BCAs have been devised as part of the research, and companies selling these products may use the said criteria. Such application will ensure the correct BCAs are used and, more specifically, used under the correct conditions. The powdery mildew (PM) fungus is often seen to develop resistance to synthetic fungicides and, therefore, alternative control measures are required. BCAs as an alternative pose less risk to the environment, workers and the consumer. A pre-trial has been conducted with a range of BCAs to see if they can control powdery mildew (PM) in a greenhouse environment on hydroponically grown cucumber (Cucumis sativus L.) plants using the variety Baccara that has only a moderate tolerance to PM. The BCAs have been compared to the control (synthetic fungicide: Bravo). Comparative work includes Coyier's model, which has been modified and adapted for these trials to determine the percentage of leaf area covered by the PM infection. Furthermore, the number of fruit harvested per treatment, kilogram yield, total mass of yield and average fruit mass is also used to determine the efficacy of the BCAs as these factors have economic significance to commercial growers. The pre-trial showed promise until the fertigation computer failed, resulting in a nutrient shortage and imbalance, confirming that BCAs alone cannot control PM. Synthetic fungicides were applied until control of PM and plant nutrition was regained. BCAs were re-introduced and used until the end of crop production. The confirmation from the pre-trial has led to the inclusion of silicon in conjunction with the BCAs in the two subsequent trials (Trials 1 & 2). Silicon was applied with the BCAs as a foliar spray on a weekly basis. In trials 1 and 2, the cucumber variety, Palladium, with a high genetic tolerance to PM is used, as this variety is suited to form part of the holistic approach used for trials 1 and 2. Trial 1 showed that treatment A, containing Streptomyces griseovirdis and Streptomyces aureofaciens, had the highest yield. Both of these are bacterial BCAs and demonstrated their adaptability to varied climatic conditions, notably when low humidity was experienced. In treatment B, Trichoderma harzianum strains, Rifai and Uppington, show the slowest rate of PM development. In trials 1 and 2, the best actual PM control was obtained by two fungal based BCAs (Trial 1, treatment C was Ampelomyces quisqualis) and (Trial 2, treatment B was Trichoderma harzianum strains, Rifai and Uppington), showing that fungal BCAs have a place for this application, but the growth-enhancing properties of bacterial based BCAs make economic sense and would make them attractive to growers. Treatment A (Streptomyces spp.) had the most number of fruit for the entire growing period and the best overall yield (kg yield) again. Two of the BCA / silicon treatments have marginally better PM control compared to that of the control (E) treatment, although not statistically significant. Treatment E (control) has the highest average fruit mass in this instance but does not have the highest yield (kg yield) when compared to treatments A and B, possibly due to the growth-enhancing properties of most of these BCAs. Therefore, most of these BCA treatments give fairly inconsistent results that vary possibly according to season, humidity and temperature, making it difficult to predict their efficacy. Using combinations or weekly alternations of these BCAs with extremes of climatic adaptation will probably be the most reliable method of obtaining consistent results. Bacterial BCAs are shown to have lower humidity requirements and produce the most consistent results in terms of fruit number, yield and fruit mass and a combination of bacterial and fungal based BCAs would possibly be the best as this would control PM and yet still have the growth enhancing properties from the bacterial based BCAs. From the research, it can be said that some BCAs in trials 1 and 2 produce results similar to that of the control in terms of percentage leaf area covered by PM and some are shown to have improved yields. Results produced from certain BCA treatments are thus equal to the control; yet provide an environmentally friendly alternative to synthetic fungicides. Silicon is listed as a beneficial element rather than an essential element; however, literature claims it to be highly effective in treating PM in cucurbits. Results from trials 1 and 2 show that control of PM is possible in most cases, when a holistic approach is used. This approach includes a cucumber variety with a high PM tolerance, optimum nutrition, cultural practices and silicon in combination with the BCAs. A complete change of management practices is necessary to implement such a BCA program. / Agriculture, Animal Health and Human Ecology / M. Tech. (Nature Conservation)