Resistance Et Évolution Des Poux Humains Pediculus Humanus

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Resistance Et Évolution Des Poux Humains Pediculus Humanus AIX-MARSEILLE UNIVERSITE ECOLE DOCTORALE DES SCIENCES DE LA VIE ET DE LA SANTE FACULTE DE MEDECINE DE MARSEILLE Unité de Recherche Microbes, Evolution, Phylogeny and Infection (MEФI) IHU-Méditerranée infection Thèse présentée pour obtenir le grade de Doctorat d’Aix-Marseille Université Spécialité Pathologie Humaine : Maladies infectieuses Mme Nadia AMANZOUGAGHENE -MEHALLA Resistance et évolution des poux humains Pediculus humanus Soutenue le 05 Juillet 2018 devant le jury : Mme le Professeur Fabienne BREGEON Présidente du jury Mr le Docteur Arezki IZRI Rapporteur Mr le Professeur Lionel ZENNER Rapporteur Mr le Docteur Oleg MEDIANNIKOV Co-directeur de thèse Direction de la Thèse : Mr le Professeur Didier RAOULT Directeur de thèse Mr le Docteur Oleg MEDIANNIKOV Co-directeur de thèse Année Universitaire 2017-2018 Remerciements Je commencerai par remercier le Professeur Didier RAOULT pour m’avoir accueilli au sein de son laboratoire et permis de réaliser ces travaux de recherche sous sa direction et ses conseils avisés. Permettez-moi de vous exprimer mon profond respect. Mes profonds remerciements au Docteur Oleg MEDIANNIKOV pour avoir dirigé ce travail. Vous m’avez donné un support scientifique incontournable et la liberté d’évoluer comme un chercheur. Merci pour m’avoir donné votre temps, votre patience, votre assistance, vos conseils et surtout votre confiance dans toutes les étapes de ce travail. Aussi grande que puisse être ma gratitude, soyez assuré qu'elle ne sera jamais à la hauteur de tous les efforts que vous avez déployé, je vous témoigne le plus profond de mes plaisirs de travailler avec vous. Mes sincères et chaleureux remerciements vont au Professeur Florence FENOLLAR, pour avoir co-dirigé ce travail. Votre aide scientifique inestimable, votre soutien moral, votre compréhension et votre gentillesse m’ont beaucoup marqué. J’espère que ce modeste travail témoigne de ma profonde reconnaissance et de ma haute considération. Je tiens à remercier très chaleureusement le Docteur vétérinaire Bernard DAVOUST pour son soutien et son aide précieuse. Trouvez ici, l'expression de ma profonde gratitude. Je remercie sincèrement Madame le Professeur Fabienne BREGEON d’avoir accepté d’examiner et de présider mon jury de thèse. Je remercie très sincèrement le Docteur Arezki IZRI et le Professeur Lionel ZENNER d’avoir accepté d’évaluer ce travail. Veuillez trouver ici l’expression de ma profonde reconnaissance. J’adresse mes vifs remerciements et ma profonde reconnaissance au Docteur Idir BITAM pour son aide et pour la confiance dont il m’a investi en me proposant de rejoindre l’IHU- méditerranée infection (ex-URMITE). Je remercie tous mes co-auteurs que je ne pourrais citer individuellement car ils sont nombreux. Mes sincères remerciements s’adressent à Jean-Michel BERENGER, Yassina BECHAH, Amira BEN AMARA, Claude NAPPEZ, Pascal WEBER et El Hadji Amadou NIANG, pour leurs conseils et leurs aides précieuses. Mes plus sincères remerciements vont à l’ensemble des étudiants, technicien(ne)s et au personnel administratif de laboratoire. Plus particulièrement Annick ABEILLE, Elsa PRUDENT, Emilie, Anne-Marie, François et mes très chères Francine VERIN et Micheline PITACCOLO pour leurs disponibilités, leurs conseils pratiques et surtout leur soutien moral et leur sourire permanent. Dédicace À mes très chers parents, ma mère et mon père, qui par leurs encouragements et leur disponibilité, m’ont poussé à persévérer et à donner le mieux de moi-même Qu’ils trouvent ici le témoignage de ma profonde gratitude. À mon très cher mari, Messaoud. Merci pour ton soutien indéfectible. Tu as toujours suivi mes études avec intérêt et tu n’as ménagé aucun effort pour ma réussite. Que l’Eternel te garde dans sa bonté. À mes frères, à mes sœurs, à mes belles-sœurs et beaux-frères pour leur soutien et bienveillance Qu'ils trouvent ici les sentiments de ma reconnaissance. À mes très chers neveux et nièces Que ce travail vous serve d'exemple pour vous surpasser. À mes beaux-parents, belles-sœurs et beau-frère pour leur gentillesse Qu'ils trouvent ici les sentiments de ma reconnaissance. À toute ma famille. Sommaire Avant-propos ...................................................................................................................... 1 Résumé ................................................................................................................................... 2 Abstract ................................................................................................................................. 4 Introduction générale ....................................................................................................... 6 Article 1. Revue : Where are we with human lice? A review of the current state of our knowledge. ........................................................................................................................................... 10 Chapitre I : Phylogénie et phylogéographie des poux anciens et contemporains ................................................................................................................... 44 Article 2 : High Ancient Genetic Diversity of Human Lice, Pediculus humanus, from Israel Reveals New Insights into the Origin of Clade B Lice. ............................................ 46 Article 3 : Mitochondrial diversity and phylogeography of Pediculus humanus, with the description of a new Amazonian Clade F ................................................................... 61 Chapitre II : Epidémiologie des poux et pathogènes associés ........................... 91 Article 4 : Head Lice of Pygmies Reveal the Presence of Relapsing Fever Borreliae in the Republic of Congo. ........................................................................................................ 93 Article 5 : Detection of several emerging bacterial pathogens in human head lice from Mali ........................................................................................................................................ 113 Article 6 : Molecular survey of Head and Body lice, Pediculus humanus, in France ............ 134 Article 7 : Detection of bacterial pathogens in clade E head lice collected from Niger's refugees in Algeria. ...................................................................................................................... 144 Article 8 : Molecular survey of head lice, Pediculus humanus capitis, in Democratic Republic of Congo ....................................................................................................................... 156 Chapitre III : Mécanismes de résistance des poux à l’ivermectine ............... 180 Article 9 : Mutations in Ivermectin-target site (GluCl) associated with field-evolved resistance of head lice recovered from Senegal. ........................................................................ 183 Article 10 : Complexin in Ivermectin resistance in body lice ................................................... 206 Conclusions et perspectives ........................................................................................ 245 Références bibliographiques ...................................................................................... 248 Annexes .............................................................................................................................. 251 Article 11 : Multiple Pathogens Including Potential New Species in Tick Vectors in Côte d'Ivoire ........................................................................................................................... 252 Article 12 : Molecular identification of microorganisms in chronic wounds, Republic of Guinea (Conakry) ...................................................................................................................... 271 Avant-propos Le format de présentation de cette thèse correspond à une recommandation de la spécialité Maladies Infectieuses et Microbiologie, à l’intérieur du Master des Sciences de la Vie et de la Santé qui dépend de l’Ecole Doctorale des Sciences de la Vie de Marseille. Le candidat est amené à respecter des règles qui lui sont imposées et qui comportent un format de thèse utilisé dans le Nord de l’Europe et qui permet un meilleur rangement que les thèses traditionnelles. Par ailleurs, la partie introduction et bibliographie est remplacée par une revue envoyée dans un journal afin de permettre une évaluation extérieure de la qualité de la revue et de permettre à l’étudiant de commencer le plus tôt possible une bibliographie exhaustive sur le domaine de cette thèse. Par ailleurs, la thèse est présentée sur article publié, accepté ou soumis associé d’un bref commentaire donnant le sens général du travail. Cette forme de présentation a paru plus en adéquation avec les exigences de la compétition internationale et permet de se concentrer sur des travaux qui bénéficieront d’une diffusion internationale. Professeur Didier RAOULT 1 / 285 Résumé Les poux de tête (Pediculus humanus capitis) et les poux de corps (P. h. humanus) sont des ectoparasites hématophages obligatoires qui ont coévolué avec les humains depuis des milliers d’années. L'analyse de l’ADN mitochondrial a classé les poux de tête en cinq clades (A-E) et a placé les poux de corps dans les clades A et D. Actuellement, les poux de corps sont reconnus comme les principaux vecteurs de maladies telles que le typhus épidémique, la fièvre des tranchées et la fièvre récurrente causées par Rickettsia prowazekii, Bartonella quintana, et Borrelia recurrentis respectivement,
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