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La Stéatose Hépatique Chez Le Canard Mulard

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La Stéatose Hépatique Chez Le Canard Mulard La stéatose hépatique chez le canard mulard : étude cinétique du métabolisme intermédiaire, stress cellulaire, autophagie et nouvelle approche par les microARNs Tracy Pioche To cite this version: Tracy Pioche. La stéatose hépatique chez le canard mulard : étude cinétique du métabolisme inter- médiaire, stress cellulaire, autophagie et nouvelle approche par les microARNs. Sciences agricoles. Université de Pau et des Pays de l’Adour, 2019. Français. NNT : 2019PAUU3043. tel-02881956 HAL Id: tel-02881956 https://tel.archives-ouvertes.fr/tel-02881956 Submitted on 26 Jun 2020 HAL is a multi-disciplinary open access L’archive ouverte pluridisciplinaire HAL, est archive for the deposit and dissemination of sci- destinée au dépôt et à la diffusion de documents entific research documents, whether they are pub- scientifiques de niveau recherche, publiés ou non, lished or not. The documents may come from émanant des établissements d’enseignement et de teaching and research institutions in France or recherche français ou étrangers, des laboratoires abroad, or from public or private research centers. publics ou privés. THÈSE UNIVERSITE DE PAU ET DES PAYS DE L’ADOUR École doctorale 211 - SCIENCES EXACTES ET LEURS APPLICATIONS Présentée et soutenue le 18 Décembre 2019 par Tracy Pioche pour obtenir le grade de docteur de l’Université de Pau et des Pays de l’Adour Spécialité : Aspects Moléculaires et Cellulaires de la Biologie La stéatose hépatique chez le canard mulard : étude cinétique du métabolisme intermédiaire, stress cellulaire, autophagie et nouvelle approche par les microARNs MEMBRES DU JURY RAPPORTEURS Bertrand PAIN Directeur de Recherche, UMRS 1208 INSERM, UCBL, INRA Hervé GUILLOU Directeur de Recherche, UMR 1331 TOXALIM INRA EXAMINATEURS Fabien SKIBA Responsable R&D Nutrition, NUTRICIA Hervé REMIGNON Professeur des Universités, UMR 1388 GenPhySE, INRA, INP- ENSAT, ENVT PRESIDENT Jean-Marc ANDRE UMR CNRS 5218, ENSC DIRECTEURS Sandrine SKIBA-CASSY Directeur de recherche, UMR 1419 INRA Karine GONTIER Maître de conférences, UMR 1419 INRA 0 1 Résumé L’objectif principal de cette thèse était de poursuivre les investigations menées sur les mécanismes sous-jacents au développement du foie gras chez le canard mulard. Ce processus appelé stéatose hépatique et induit par le gavage nécessite d’être davantage étudié afin de contribuer à l’optimisation du gavage. Une étude cinétique de l’expression des gènes en lien avec la stéatose hépatique, le métabolisme intermédiaire et le stress cellulaire chez le canard mulard a permis de mettre en évidence, outre la forte mobilisation des métabolismes glucidique et lipidique, une induction significative de la voie de signalisation de l’insuline mTOR et l’activation de mécanismes de protection cellulaire au sein des hépatocytes leur permettant de lutter contre l’apport accru d’amidon apporté par le gavage. Parmi ces différents mécanismes protecteurs, l’autophagie a fait l’objet d’une étude poussée chez des canards gavés dont la stéatose a été à peine induite comparés à des canards non gavés. Les mesures de flux autophagique (LC3-II) ont montré que le gavage induisait une inhibition du processus autophagique dans les hépatocytes de canards mulards dès la phase précoce du développement de la stéatose hépatique. Cette même étude a permis de constater que les mécanismes associés au stress cellulaires tels que le stress oxydant, le stress du réticulum endoplasmique ou encore l’apoptose ne se sont pas induits dès le début du développement de la stéatose hépatique mais bien plus tardivement. Il en est de même pour l’impact du gavage sur la voie mTOR. Par ailleurs, l’étude cinétique a également permis de mettre en évidence l’induction significative du métabolisme du cholestérol corrélé au gain de poids de foie des canards au cours du gavage. Ces résultats ont suscité l’intérêt de rechercher des biomarqueurs plasmatiques du développement de la stéatose hépatique en lien avec le métabolisme du cholestérol parmi les microARNs. Ainsi, le miRNome plasmatique du canard a été séquencé et a permis de définir les microARNs les plus différentiellement exprimés dans le plasma des canards mulards mais également de déceler des microARNs (miR-122, miR-107 et miR-194) candidats comme potentiels biomarqueurs utilisables pour le suivi du développement du foie gras. L’ensemble de ces travaux contribuent à l’avancée de la recherche sur la compréhension des mécanismes de l’établissement de la stéatose hépatique chez le canard mulard. Mots clés : stéatose hépatique, canard mulard, gavage, métabolismes, autophagie, microARNs 2 Abstract The main objective of this thesis was to continue the investigations carried out on the mechanisms underlying the development of foie gras in mule ducks. This process, known as hepatic steatosis and induced by overfeeding, needs to be further studied in order to contribute to overfeeding optimization. A kinetic study of the expression of genes related to hepatic steatosis, intermediate metabolism and cellular stress in mule ducks highlighted firstly the strong mobilization of carbohydrate and lipid metabolisms. Secondly a significant induction of the insulin / mTOR signalling pathway and the activation of various cellular protection mechanisms in hepatocytes were observed allowing them to fight against the increased intake of starch brought by the overfeeding. Among these are various protective mechanisms, autophagy has been the subject of extensive study in overfed ducks whose steatosis has been barely induced compared to non-overfed ducks. Autophagic flux measurements (LC3-II) showed that overfeeding induced inhibition of the autophagic process in mule ducks’ hepatocytes from the early stage of hepatic steatosis. This same study found that the mechanisms associated with cellular stress such as oxidative stress, endoplasmic reticulum stress or apoptosis did not occur early in the development of hepatic steatosis but much later. The same is true for the impact of overfeeding on mTOR signalling pathway. In addition, the kinetic study also revealed the significant induction of cholesterol metabolism correlated with the liver weight gain of ducks during overfeeding. These results aroused the interest of investigating plasma biomarkers for the development of hepatic steatosis related to cholesterol metabolism among microRNAs. Thus, duck's plasmatic miRNome was sequenced and it allowed to define the most differentially expressed microRNAs in the plasma of mule ducks but also to detect microRNAs (miR-122, miR-107 and mi-R194) as potential biomarkers that could be used for monitoring the development of fatty liver. All of this work contributes to the advancement of research on understanding the mechanisms of establishment of hepatic steatosis in mule ducks. Key words: hepatic steatosis, mule duck, overfeeding, metabolism, autophagy, microRNAs 3 Sommaire Travaux issus de la thèse…………………………………………………………………….....7 Abréviations……………………………………………………………………………………9 Figures………………………………………………………………………………….…..…13 Tableaux…………………………………………………………………………………..…..15 Introduction générale…………………………………………………………………......…..16 Etude bibliographique…………………………………………………………...………...….20 1. Modèle d’étude : le canard mulard ................................................................................... 22 2. Le gavage .......................................................................................................................... 23 3. Le métabolisme des oiseaux ............................................................................................. 25 3.1 Métabolisme glucidique ............................................................................................ 27 3.1.1 Digestion, absorption du glucose ....................................................................... 27 3.1.2 Utilisation du glucose ......................................................................................... 27 3.2 Métabolisme lipidique ............................................................................................... 30 3.2.1 Lipogenèse de novo ............................................................................................ 30 3.2.2 Devenir des acides gras néo-synthétisés ............................................................ 32 3.2.3 Transport et exportation ..................................................................................... 32 3.2.4 Captage ............................................................................................................... 34 3.2.5 La β-oxydation ................................................................................................... 36 3.2.6 Métabolisme du cholestérol ............................................................................... 37 3.3 Régulation des métabolismes intermédiaires............................................................. 40 3.3.1 L’insuline ........................................................................................................... 40 3.3.2 La protéine Akt ................................................................................................... 43 3.3.3 La voie TOR et ses effecteurs ............................................................................ 45 3.3.4 Les acides aminés ............................................................................................... 47 4. La stéatose hépatique ........................................................................................................ 50 4.1 Chez les mammifères ................................................................................................
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