Approches Evolutive Et Mecanistique

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Approches Evolutive Et Mecanistique THESE DE DOCTORAT DE L’ETABLISSEMENT UNIVERSITE BOURGOGNE FRANCHE-COMTE PREPAREE A L’UNITE MIXTE DE RECHERCHE CNRS 6282 BIOGEOSCIENCES Ecole doctorale n°554 Environnement, Santé Doctorat des Sciences de la Vie Spécialité Ecologie Evolutive Par Fayard Marion _______________________________________________________________________________________ ANXIETE ET MANIPULATION PARASITAIRE CHEZ UN INVERTEBRE AQUATIQUE : APPROCHES EVOLUTIVE ET MECANISTIQUE Thèse présentée et soutenue à Dijon, le 28 Août 2020 Composition du Jury : Jean-Nicolas Beisel, Professeur, ENGEES, Université de Strasbourg Rapporteur Anne-Sophie Darmaillacq, Maître de Conférences, Université de Caen Rapporteure Jean-François Ferveur, Directeur de recherches CNRS, Université de Bourgogne Franche-Comté Examinateur Vincent Médoc, Maître de Conférences, Université de Saint-Etienne Examinateur Marie-Jeanne Perrot-Minnot, Maître de Conférences, Université de Bourgogne Franche-Comté Directrice Thierry Rigaud, Directeur de recherches CNRS, Université de Bourgogne Franche-Comté Examinateur - Président Titre : Anxiété et manipulation parasitaire chez un invertébré aquatique : approches évolutive et mécanistque Mots clés : acanthocéphale, amphipode, comportement, état émotionnel, manipulation parasitaire, prédation Les parasites à transmission trophique sont connus pour les de transmission, est faible. Chez les individus sains, nous avons mis changements phénotypiques qu’ils induisent chez leurs hôtes. Ces en évidence, par une approche corrélationnelle, une variabilité changements sont supposés favoriser la transmission des parasites interpopulationnelle de l’intensité des comportements antiprédateur vers un hôte définitif à travers la prédation de l’hôte intermédiaire. en lien avec le risque local de prédation, et la densité de gammares Ce phénomène de « manipulation parasitaire » a longtemps été vu conspéficiques. Nous avons également montré une flexibilité de la comme un trait adaptatif des parasites. La manipulation reposant réponse au stimulus de prédation chez des individus provenant de sur des interactions proie-prédateur, il est nécessaire de comprendre populations où le risque de prédation était élevé, la réponse comment les comportements antiprédateur sont modulés par des augmentant avec l’intensité du stimulus. En revanche, les individus facteurs exogènes (pression de prédation) et endogènes (infection provenant de populations à faible risque de prédation semblent parasitaire, état émotionnel, …) aux individus. Au cours de cette montrer une réponse relativement forte indépendamment de thèse, nous avons tenté d'approfondir la compréhension de ce l’intensité du stimulus, ce qui suggère une hypersensibilité. Ces phénomène, chez les crustacés amphipodes, en répondant à études corrélationnelles, portant sur l'analyse de la variabilité plusieurs questions : (1) quelle est l'étendue de la interpopulationnelle selon les pressions de prédation locales, nous multidimensionnalité de la manipulation parasitaire par les ont amenés à supposer que ces différentes stratégies seraient Acanthocéphales, quantifiée au travers d'une méta-analyse ; (2) intimement liées à l’expérience d’un stress chronique de la l'amplitude des changements de comportements antiprédateur prédation. Nous suggérons alors l’existence d’un état de long-terme varie-t-elle selon le contexte local de prédation ? ; (3) les de type anxiété qui pourrait être la résultante de la répétition comportements antiprédateur présentent-ils une flexibilité en lien d’épisodes de court-terme de peur. Nous avons effectué un premier avec le contexte local de prédation ? ; (4) le parasite "exploite" t-il pas en montrant expérimentalement l’existence de comportements la flexibilité comportementale des gammares sains, notamment en de peur et de type anxiété chez G. fossarum. L’ensemble de ces lien avec leur régulation émotionnelle de type anxiété ?. Au niveau travaux a démontré la place centrale des interactions proie- interspécifique à l'échelle du phylum des Acanthocéphales, nous prédateur dans l’étude de la manipulation parasitaire. Plus avons mis en évidence une altération marquée des comportements précisément, nous avons mis en évidence une variabilité et une en lien avec la défense antiprédateur des hôtes (changement de modulation complexe des comportements antiprédateur des hôtes microhabitat, protection et réponse à des stimuli). Au niveau du en lien avec le contexte local de prédation, et qui pourraient trouver couple hôte-parasite Gammarus fossarum, Pomphorhynchus racine dans un état émotionnel lié au stress chronique de la tereticollis, nous avons montré que l’intensité des changements de prédation. Ces travaux ouvrent alors quelques pistes à comportements antiprédateur induit par l'infection (phototaxie et investiguer telles que les mécanismes sous-jacents de cet état et utilisation du refuge) présentait une variabilité interpopulationnelle, l’éventuel rôle des parasites dans la modulation de cet état de type en lien avec le risque de prédation : la manipulation semble d’autant anxiété qui viendrait modifier l’expression des comportements plus forte que la pression de prédation locale, i.e. les opportunités antiprédateur. Title : Anxiety and parasite manipulation in an aquatic invertebrate : evolutive and mechanistic approaches Keywords : acanthocephala, amphipoda, behaviour, emotional state, parasite manipulation, predation Trophically transmitted parasites induce changes in their host’s transmission opportunities, is low. With a correlational approach, phenotype. These changes are supposed to increase transmission we also evidenced variation in the magnitude of antipredatory probability to definitive hosts through the predation of intermediate behaviours according to local predation risk and conspecifics hosts. This phenomenon is known as ‘parasite manipulation’ has density, in uninfected individuals. In addition, we emphasized been hypothesized to be an adaptive trait of parasites for a long flexibility of behavioural responses to predator cues: individuals time. As manipulation involves predator-prey interactions, it is from populations experiencing high predation risk exhibited therefore necessary to understand how antipredatory behaviours increased responses as predator cues concentrations increased. In are modulated by exogenous (predation pressure) and endogenous contrast, individuals from populations experiencing low predation (infection, emotional state) factors. We tried to go into this risk appeared to exhibit strong responses independent of predator phenomenon in depth, in amphipods, by responding to several cues concentrations, suggesting hypersensitivity. We supposed that questions : (1) what is the extent of the multidimensionality in these strategies would be closely related to the exposure to chronic parasite manipulation by Acanthocephalan, quantified through a predation risk. We therefore suggest the existence of an anxiety- meta-analysis ; (2) is there variation in the magnitude of like state that could result from the repetition of acute stress, i.e. antipredatory behaviours according to local predation risk? (3) are fear, episodes. We made a first step through an experimental antipredatory behaviours flexible with respect to local predation approach, evidencing the existence of both anxiety-like and fear risk? (4) Do parasites exploit behavioural flexibility in uninfected behaviours in G. fossarum. Overall, these works pointed out the individuals, in relation to an emotional state such as anxiety-like importance of predator-prey interactions in the study of parasite state? Within the Acanthocephala phylum, we evidenced notable manipulation. More particularly, we evidenced variation and changes, more particularly of host antipredatory behaviours complex modulation of host antipredatory behaviours in (microhabitat changes, protection and responses to stimuli). accordance with local predation risk, and which may be closely Considering Gammarus fossarum, Pomphorhynchus tereticollis related to an emotional state stemming from chronic predation host-parasite couple, we showed that there was variation in the stress. We therefore suggest some future directions to investigate, magnitude of antipredatory behavioural changes induced by such as the underlying mechanisms of anxiety-like state, and the infection (phototaxy and refuge use), with respect to local predation role of parasite in the modulation of this state that would modify risk : host manipulation seemed as strong as predation risk, i.e. the expression of antipredatory behaviours. Université Bourgogne Franche-Comté 32, avenue de l’Observatoire 25000 Besançon TABLE DES MATIERES INTRODUCTION GENERALE …………………………………………………………… p.1 Contexte de la manipulation parasitaire …………………………………………. p.2 Le comportement : dimension phénotypique phare ? ……………………………. p.6 Modèles biologiques : les acanthocéphales et gammares ………………………... p.8 Objectifs de la thèse………………………………………………………………. p.15 PARTIE 1 – BILAN CRITIQUE ET SUR LA MANIPULATION PARASITAIRE ……………… p.22 Chapitre 1 – Magnitude and direction of parasite-induced phenotypic alterations ……………. p.23 Discussion-Transition ………………………………………………………………………….. p.75 Variation liée au parasite ou à l’hôte ………………………………………………………..... p.75 Influence des facteurs environnementaux ……………………………………………………. p.77 PARTIE 2 – VARIATION INTERPOPULATIONNELLE DE LA MANIPULATION ………….. p.80 Chapitre 2 – Interpopulation variation in the intensity of host manipulation …………..……… p.81 Transition ………………………………………………………………………… p.117 Chapitre
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