these-Navin-last printing ver number in middle - Copiethesesups.ups-tlse.fr/3174/1/2016TOU30052.pdf · Je tiens à remercier infiniment à Madame le docteur Béatrice Cousin qui a

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REMERCIMENTS

Voici mes 3 années de thèse qui s’achèvent. C’est maintenant le moment de rédiger une partie

très importante: les remerciements. La rédaction de cette partie a commencé dans mon coeur

dès les premiers jours de mon arrivée en France, séparée de mon aimable pays, le Cambodge.

J’adresse tout d’abord ma très sincère reconnaissance à mon directeur de thèse, Monsieur le

Professeur Rémy Burcelin (INSERM 1048), qui m’a accueillie chaleureusement au sein de

son équipe et m’a guidée tout au long de ce travail. C’est dans ce labo, que j’ai découvert de

nouvelles techniques et où j’ai obtenu les premiers résultats. Je le remercie profondément

pour ses conseils éclairés, ses corrections précieuses pour nos articles et pour cette thèse. Je

garderai beaucoup de considération pour sa force de travail.

Je tiens également à exprimer ma reconnaissance à mon co-directeur de thèse Monsieur le

Docteur Eric Serée (INSERM 1062 / INRA 1260) qui m’a invitée à travailler dans son

équipe et m’a proposé ce sujet de thèse très intéressant. Du fond du coeur, je voudrais lui

adresser, mes plus sincères remerciements. Je le remercie également pour sa confiance, son

soutien, et pour sa présence dans tous les moments difficiles de cette thèse.

Je n’aurais pu avoir la possibilité de réaliser ce travail de thèse sans mon encadrant Monsieur

le Professeur Serge Champion (CNRS 7263 – IMBE) qui m’a partagé ses connaissances et

ses compétences en Biochimie et Biologie Moléculaire et qui m’a donné de nombreux

conseils pour mener à bien ce travail. Je le remercie sincèrement pour son aide précieuse dans

la préparation de cette revue de thèse ainsi que des différentes communications réalisées lors

de cette thèse.

Je tiens à remercier également à Monsieur le Professeur Yves Barra qui m’a accepté dans

son laboratoire. Je le remercie tout particulièrement pour sa gentillesse, son soutien et ses

conseils précieux.

Je voudrais remercier Madame le Docteur Catherine Riva, Madame le Professeur Evelyne

Ollivier et Monsieur le Professeur Philippe Piccerelle de m’avoir fait l’honneur d’être

rapporteurs de cette thèse. Je suis très honorée de leur investissement dans la lecture et les

corrections de cette thèse. Je vous prie, Mesdames, Monsieur, de trouver ici le témoignage de

ma reconnaissance et de mon profond respect.

#!

Je tiens à remercier infiniment à Madame le docteur Béatrice Cousin qui a accepté d’être

une des membres de mon comité de thèse. Je la remercie pour sa participation et ses conseils

précieux.

Tous mes remerciements vont également à Madame le Professeur Marie-Christine Alessi,

la Directrice de l’UMR INRA 1260, pour m’avoir accueillie dans son unité et m’avoir assuré

des conditions de travail de bonne qualité. Je vous prie, Madame, de trouver ici l’expression

de ma reconnaissance.

Les résultats que j’ai la chance de présenter dans cette thèse sont le fruit d’un travail

d’équipes. Je voudrais adresser mes remerciements sincères à tous les personnels et collègues

des équipes ci-dessous:

- L’équipe 2 "Facteurs de Risques Intestinaux, Diabète, Dyslipidémie", INSERM

U1048, I2MC, Institut de Recherche sur les Maladies Métaboliques et

Cardiovasculaires de l'Hôpital Rangueil, Toulouse.

- L’équipe IMBE de Mutagénèse environnementale de l’UMR CNRS 7263, Faculté de

Pharmacie, Aix Marseille Université.

- L’équipe 2 "Micronutriments, Tissu adipeux et Résistance à l’insuline", l’UMR

INSERM 1062 / INRA 1260, "Nutrition Obésité et Risque Thrombotique", Faculté de

Médecine, Aix Marseille Université.

- L’équipe de la Zootechnie, I2MC, Institut de Recherche sur les Maladies

Métaboliques et Cardiovasculaires de l'Hôpital Rangueil, Toulouse.

- La plateforme d’imaginerie, I2MC, Institut de Recherche sur les Maladies

Métaboliques et Cardiovasculaires de l'Hôpital Rangueil, Toulouse.

- Le plateau de Génomique Get-Purpan, IFR150, Institut Fédératif de Recherche Bio-

Médicale de Toulouse (IFR-BMT) de l’Hôpital Purpan, Toulouse.

- L’équipe de Vaiomer, Labege, Toulouse.

Merci à toutes celles et ceux qui m’ont acceptée dans une ambiance agréable, merci pour leur

disponibilité, leur gentillesse et leur soutien permanent souvent technique, parfois moral:

Pascale Loubières: je te remercie infiniment pour ton accueil chaleureux au laboratoire. Je ne

me sens jamais seule avec toi. Merci énormément d’être à mes côtés pour m’écouter, me

rassurer et partager des bons moments. J’aime te raconter des choses et discuter avec toi et

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partager des connaissances sur le monde des bactéries. Je remercie également à toute ta

famille pour leur accueil chaleureux pendant mon séjour à la montagne. J’ai passé une

merveille randonnée avec eux.

Jean-Charles Martin: je vous remercie beaucoup pour les données GC-MS.

Patricia Bremond: sans toi, je n’aurai pas pu apprendre et réussir toutes les cultures de

cellules qui ont été nécessaires. Je te remercie beaucoup.

Vincent Azalbert: merci de m’avoir appris à faire le DGGE qui est une technique

compliquée à réaliser. Je n’aurai pas pu le réussir sans ton aide.

Céline Pomié et Estelle Grasset: je vous remercie de m’avoir partagé vos échantillons pour

mener à bien mes expériences sur les cytochromes.

François Tercé: merci beaucoup pour ton soutien chaleureux et tes conseils précieux

concernant les expériences sur les lipides.

Vincent Blasco: je te remercie de m’avoir appris à faire l’analyse des résultats.

Céline Garret: merci de m’avoir appris les extractions d’ARN.

Matteo Serino: merci beaucoup pour tes conseils et pour m’avoir appris à réaliser

l’extraction d’ADN.

Aurélie Waget: je te remercie pour m’avoir appris les manipulations des souris.

Jean François-Landrier et Frank Peiretti: merci pour votre aide technique.

Mes remerciements vont également à Monsieur le Professeur Philippe Valet et son équipe

du service de l’Ecole doctorale de Biologies Santé et Biotechnologies pour leur sympathie et

leur aide pour surmonter les processus administratifs lors de la soutenance de cette thèse.

Je tiens de remercier l’Université Toulouse III – Paul Sabatier et le coordinateur du Projet

Techno II – Erasmus Mundus pour m’avoir octroyé une bourse afin de réaliser cette thèse. Je

remercie infiniment Monsieur Jean-Michel Baleynaud, Monsieur le Professeur Sovan

Lek, Mademoiselle Laura Verheyleweghen et leur équipe du Projet Techno II.

Je remercie énormément Madame Sophie BOCHET et son équipe du Service d’accueil des

étudiants étrangers du CROUS Aix-Marseille pour leur sympathie et pour les efforts qu’ils

ont consacré à me trouver un logement à Marseille.

Je voudrais adresser mes sincères remerciements au Doyen de la Faculté de Pharmacie du

Cambodge pour son soutien et à Monsieur le Docteur Rothmony Eang pour son aide

administrative pour ma demande de bourse.

%!

Je tiens à remercier tout particulièrement Monsieur Hot Bun et sa famille et Monsieur

Chhan Siv et son épouse pour leurs conseils précieux, leur soutien et leurs encouragements.

Je voudrais remercier également à l’AEPK (Amicale des étudiants en Pharmacie et des

Pharmaciens Khmer) qui aide techniquement et financièrement les étudiants cambodgiens.

Je remercie spécialement mes deux très bonnes amies, Hanna Ilchmann et Mardy Long qui

sont toujours à mes côtés. Hanna, je te remercie pour m’avoir partagé ta chambre durant

chacun de mes séjours à Toulouse. Merci à toutes les deux pour votre soutien et votre

encouragement. Pour moi, vous êtes comme mes sœurs.

Mes 3 années de thèse en France ont été marquées par l’enthousiasme, la joie, les surprises et

aussi quelquefois les doutes, l’angoisse, la démotivation…Je voudrais remercier tous mes

amis cambodgiens et étrangers d’avoir partagé avec moi ces sentiments. Je vous remercie

profondément pour votre présence, votre patience, votre soutien et votre encouragement.

Du fond du coeur, je tiens à remercier toute ma famille: Papa, Maman, ma grande mère, mes

sœurs, mon petit frère, mon parrain et ma marraine, mon beau frère, mes tantes, mes oncles,

mes cousins, cousines, mon neveu Lin Moon …. Votre amour, votre soutien, vos

encouragements permanents m’ont permis d’aller jusqu’au bout de ce long chemin.

Enfin, merci à toi, mon prince, Sopheap Oeng, de m’avoir assuré que tu es toujours là, pour

moi. Ces années passées en France ont été, pour nous deux, une épreuve à surmonter afin

d’atteindre une relation éternelle. Merci de m’aimer si fort tous les jours.

!!!!!!!!!!!!!

&!

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"%!

LISTE DES TABLEAUX

!!-ABJ>AR!"f!.NRKXDEF>!?R!?@ABCD>!QNHNOGH@TR>!//////////////////////////////////////////////////////////////////////////////////!Y"!

-ABJ>AR!#f!;CH>K!>D!JNSRK!J>K!FJRK!IGFJ@TRGK!3-#!KRKS>FD@B@J@DG!/////////////////////////////////////////////////!Y#!

-ABJ>AR!$f!*NHD>HR!?>!'.6!?AHK!?@LLGI>HD>K!KNRIS>K!//////////////////////////////////////////////////////////////////////!=%!

!

!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!

"&!

LISTE DES FIGURES

!

!"#$%&'()'6N@>K!?>!K@OHAJ@KAD@NH!*BJh*0<!?>!+'!/////////////////////////////////////////////////////////////////////////////////!##!

!"#$%&'*)'6N@>K!?>!K@OHAJ@KAD@NH!<+$j!>D!)0<!e@HAK>!?>!+'.!////////////////////////////////////////////////////////!#$!

!"#$%&'+)'(LL>D!?P@HKRJ@H>!KRI!JPMNQGNKDAK@>!OJRS@?@TR>!/////////////////////////////////////////////////////////////////!#&!

!"#$%&',)!2>K!SNHKGTR>HS>K!FND>HD@>JJ>K!?>K!FMNKFMNIEJAD@NHK!?>K!FINDG@H>K!+'.!KRI!J>K!

.>Ih-MI!/////////////////////////////////////////////////////////////////////////////////////////////////////////////////////////////////////////////!#=!

!"#$%&'-)!.DIRSDRI>!>D!FAID>HA@I>K!?k@HD>IASD@NH!?>K!+'./!//////////////////////////////////////////////////////////////!#5!

!"#$%&'.)!2k@HM@B@D@NH!?>!JA!WN@>!?>!K@OHAJ@KAD@NH!?>!Jk@HKRJ@H>!FAI!J>!.7*.X"!>D!.7*.X$/!/!$U!

!"#$%&'/)!2>K!WN@>K!@QFJ@TRG>K!?AHK!JP@HKRJ@HNXI>K@KDAHS>!?AHK!J>!QRKSJ>/!///////////////////////////!$8!

!"#$%&'0)!2>K!WN@>K!?R!KDI>KK!?R!IGD@SRJRQ!>H?NFJAKQ@TR>!SNH?R@KAHD!V!RH>!

@HLJAQQAD@NH/!/////////////////////////////////////////////////////////////////////////////////////////////////////////////////////////////////!%"!

!"#$%&'1)!2P@HLJAQQAD@NH!>D!JA!IGK@KDAHS>!V!JP@HKRJ@H>f!WN@>K!?>!K@OHAJ@KAD@NH!@QFJ@TRG>K/%$!

!"#$%&'(2)!2A!IGFNHK>!@QQRH@DA@I>!?AHK!J>!D@KKR!A?@F>R[!NBCK>K/!///////////////////////////////////////////!&U!

!"#$%&'(()!2PGWNJRD@NH!?>!S>JJRJ>K!Z!?AHK!JA!SNQF>HKAD@NH!?>!JP@HKRJ@HNXIGK@KDAHS>/!///////!&Y!

!"#$%&'(*)!2A!LJNI>!@HD>KD@HAJ>!>D!J>!QGDABNJ@KQ>!?>!JkM\D>/!/////////////////////////////////////////////////////////!=#!

!"#$%&'(+)!2>K!?>R[!@KNQCI>K!?R!'.6!////////////////////////////////////////////////////////////////////////////////////////////////////!=&!

!"#$%&'(,)!2A!WN@>!QGDABNJ@TR>!?R!'>KWGIADINJ!?AHK!JP@HD>KD@H>!FAI!J>K!>HcEQ>K!4;-K/!=Y!

!"#$%&'(-)!2A!WN@>!QGDABNJ@TR>!?R!'>KWGIADINJ!?AHK!J>!LN@>!FAI!J>K!>HcEQ>K!.42-K/!//////!==!

!"#$%&'(.)!2>K!>LL>DK!B@NJNO@TR>K!?R!'>KWGIADINJ/!////////////////////////////////////////////////////////////////////////////!=5!

!"#$%&'(/)!2>K!S@BJ>K!B@NJNO@TR>K!QNJGSRJA@I>K!KRFFNKG>K!?R!'>KWGIADINJ!?AHK!J>K!

FINS>KKRK!AHD@X@HLJAQQADN@I>K!>D!AHD@XN[E?AHD>K/!////////////////////////////////////////////////////////!5U!

!"#$%&'(0)!2>K!?>R[!@KNQCI>K!?>!*RISRQ@H>/!/////////////////////////////////////////////////////////////////////////////////////!5$!

!"#$%&'(1)!2>K!KDIRSDRI>K!?>!JA!*RISRQ@H>!>D!K>K!QGDABNJ@D>K!///////////////////////////////////////////////////!5&!

!"#$%&'*2)!)N?RJAD@NH!FAI!JA!*RISRQ@H>!?>!JP>[FI>KK@NH!?>!?@LLGI>HD>K!QNJGSRJ>K!?>!

K@OHAJ@KAD@NH!S>JJRJA@I>/!//////////////////////////////////////////////////////////////////////////////////////////////////////////////!5Y!

"Y!

!"#$%&'*()!)GSAH@KQ>!?PASD@NH!?>!JA!*RISRQ@H>!KRI!JA!WN@>!?>!K@OHAJ@KAD@NH!?>!,:Xp1/!/!55!

!"#$%&'**)!([FI>KK@NH!?>!*EFK!?AHK!J>!SNJNH/!/////////////////////////////////////////////////////////////////////////////////!"&8!

!"#$%&'*+)!([FI>KK@NH!?>!*EFK!?AHK!JP@JGNH/!/////////////////////////////////////////////////////////////////////////////////////!"YU!

!"#$%&'*,)!([FI>KK@NH!?>!*EFK!?AHK!J>!LN@>/!/////////////////////////////////////////////////////////////////////////////////////!"Y"!

!"#$%&'*-)!([FI>KK@NH!?>!*EF#S&&!>D!*EF#>"!?AHK!?@LLGI>HDK!DEF>K!?>!KNRI@K/!///////////////!"Y#!

!"#$%&'*.)!.@D>K!?PASD@NH!FNKK@BJ>K!?R!`:3!>D!?>K!FNJEFMGHNJK!//////////////////////////////////////////////!"Y&!

!"#$%&'*/)!ÈFNDMCK>!SNHS>IHAHD!J>K!K@D>K!?PASD@NH!FNKK@BJ>K!?R!`:3!>D!?>K!FNJEFMGHNJK!

>D!J>RIK!I>JAD@NHK!M@GIAISM@TR>K/!/////////////////////////////////////////////////////////////////////////////////////////!"=$!

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"5!

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"8!

34 5&'6"789:&'6&':;<&'*'

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!!!!!!!!!!!!!!!!!!!!!!!!

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8#!

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!!!!!!!!!!!

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8&!

Résultats !!!!!!!!!!!!!!!!!!!!!

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Resveratrol improves glycemia and modulates microbiota dysbiosis,

metabolome and inflammation

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Discussion et

Perspectives !!!!!!!!!!!!!!!!!!!!

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FAI! RH! IGO@Q>! I@SM>! >H! OIA@KK>K! ]`:3â! DAH?@K! TRPRH! ARDI>! FNJEFMGHNJa! JA! *RI! H>!

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!!!!!!!!!!!!!!!!!!!!!!

#"U!

Annexes !!!!!!!!!!!!!!!!!!!!!!!

#""!

(4 K>A$@:7:A'6$':7$\'6&'HDE@&A:>%E@'&:'6&':%"#@;H>%"6&A'<@7AG7:"I$&A'!!

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AR!H@W>AR!FJAKQAD@TR>!?R!DAR[!?>!DI@OJESGI@?>K!TR>JTR>!KN@D!J>!DIA@D>Q>HD/!

!!

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55 èmes Journées Nationales de Diététique et Nutrition

LE RESVÉRATROL AU SERVICE DU DIABÈTE

Dr N. SRENG1, Dr E. SERÉE1, Pr S CHAMPION2

1INRA, UMR1260 / 1062 INSERM / AMU ; Nutrition, Obésité et risque thrombotique, 27 boulevard Jean-Moulin, Marseille F-13385, France

2IMBE - UMR CNRS 7263 / IRD 237, Mutagenèse Environnementale, Aix-Marseille Université, Faculté de Pharmacie 27 Bvd Jean Moulin, Marseille F-13385,

France

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NOUVEAUX TRAITEMENTS ET PERSPECTIVES DANS LE DIABÈTE DE TYPE 2

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INTRODUCTION ET ÉTIOLOGIE DU DIABÈTE DE TYPE 2

Le diabète de type 2 (DT2) est à ce jour considéré comme une pandémie selon l’Organisation Mondiale de la Santé. Le nombre de personnes diabétiques dans le monde est actuellement estimé à environ 250 millions, et devrait atteindre près de 380 millions en 2025, ce qui représente 7,1 % de la population adulte âgée de 20 à 80 ans. En Europe, une croissance de 16 % de la maladie est prévue d’ici à 2025 [1, 2]. Si le diabète est une pathologie qui préoccupe nos sociétés, il faut tenir compte des pathologies associées. En effet, les complications cardiaques, vasculaires, rénales et ophtalmologiques, augmentent également le coût de la prise en charge du diabète et constituent actuellement, en plus d’un problème de santé publique, un risque économique pour l’ensemble des pays de l’ouest. Les causes du développement DT2 sont multiples, à la fois génétique, environnementale et sociale. La sédentarité associée à un enrichissement de la proportion de graisses dans les régimes alimentaires est un facteur aggravant qui, combiné au patrimoine génétique, est sûrement responsable de l’épidémie. Parallèlement à cela, nous sommes victimes d’une diminution de la consommation en fibres végétales qui sont impliquées dans la digestion et l’équilibre alimentaire par l’apport de micronutriments (vitamines, sels minéraux, etc.). Un autre rôle fondamental joué par ces fibres végétales concerne leur fonction de substrat à l’égard de notre flore intestinale. La diminution de leur proportion engendre un changement qualitatif de la flore intestinale et par voie de conséquence une altération du métabolisme énergétique [3-13] . Les conséquences physiopathologiques de ce déséquilibre énergétique se traduisent d’une part par une diminution de l’action de l’insuline i.e. l’insulinorésistance et d’autre part par une augmentation de la masse adipeuse. Ces deux conséquences fonctionnelles insulinorésistance et adipogénèse sont reliées entre elles via un processus inflammatoire de bas niveau qui apparaît progressivement et qu’on dénomme inflammation métabolique [3]. En utilisant des modèles animaux qui développent des maladies métaboliques tels que des souris nourries par un régime riche en graisse (> 40 % p/p) pendant plusieurs mois, on observe une augmentation de la sécrétion des cytokines inflammatoires et de chimiokines. Ces cytokines, notamment le TNF- , l’IL-1, et l’IL-6 [4, 5], interfèrent avec la signalisation générée par le récepteur à l’insuline. Ceci se traduit par une diminution de l’utilisation du glucose par les muscles et une levée d’inhibition de la production hépatique de glucose. Il s’en suit une augmentation de la glycémie puis l’apparition d’un hyperinsulinisme et d’une hyperglycémie ce qui induit la redistribution des flux métaboliques notamment vers les organes de stockage tel que le tissu adipeux et le foie. Une autre voie récemment mise en évidence montre le rôle que peuvent avoir des changements de la flore bactérienne, en liaison avec l’alimentation dans l’installation des maladies métaboliques notamment l’obésité [7, 11] et le DT2 [3-6]. Il apparaît que la modification de la flore intestinale semble

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être une des causes de l’obésité. En effet, un régime riche en graisse favorise, d’une part, l’émergence de bactéries Gram - qui seraient à l’origine de l’installation de l’inflammation métabolique et d’autre part, diminue le nombre de germes Gram + [3-5, 7-10]. Des études ont montré que le genre Bifidobacterium (Gram +) est inversement corrélé à l’incidence du DT2 et de l’obésité chez des souris soumises à un régime riche en graisse [3-5]. L’utilisation de fibres alimentaires induit une augmentation du nombre de bactéries à Gram + chez les souris nourries avec un régime riche en graisses, et parallèlement, on constate une diminution de l’inflammation et un retard dans l’installation de l’insulinorésistance [14].La thérapie antidiabétique n’est pas à ce jour efficace. En effet, les molécules actuellement sur le marché ne donnent pas les résultats escomptés et il est difficile de régler avec les traitements disponibles la complexité du désordre métabolique. Il est alors urgent de proposer de nouvelles stratégies thérapeutiques. Une alternative serait de prévenir l’apparition de la maladie chez les personnes à risque et/ou d’accompagner le traitement du diabète déclaré par la prise de compléments alimentaires. Les approches nutritionnelles et ‘alicamenteuses’ sont ainsi considérées comme bien tolérées. De nombreuses données, revues ci-dessous, suggèrent que le resvératrol (RVR) pourrait correspondre à une nouvelle stratégie thérapeutique et ‘alicamenteuse’. Son mécanisme d’action serait basé sur des effets à la fois antioxydant et anti-inflammatoire [15,16].

LE RESVÉRATROL : MOLÉCULE DE PRÉVENTION ET/OU THÉRAPEUTIQUE

Le RVR, ou 3,5,4’ transtrihydroxytilbene (figure 1), est un polyphénol naturel que l’on trouve dans de nombreuses espèces végétales dont le raisin noir, les cacahuètes, le pin et la renouée chinoise (Polygonum Cuspidatum), sont les principales sources commerciales. Le RVR est une molécule qui est synthétisée par les plantes en réponse à un stress par exposition à la lumière ultra-violette ou une attaque fongique [17, 18]. Le RVR est étudié depuis de nombreuses années et il possède de nombreuses propriétés pharmacologiques.

Le resvératrol : un antagoniste compétitif du Récepteur des Arylhydrocarbures (AhR)

Le Récepteur des Arylhydrocarbures (AhR) est un facteur de transcription dont les ligands naturels endogènes ne sont pas bien caractérisés. Le récepteur AhR est impliqué dans la régulation de l’expression de nombreux gènes lorsqu’il lie divers polluants environnementaux tels que les hydrocarbures aromatiques du tabac et des gaz d’échappement, les dioxines (dont la TCDD, Dioxine de Seveso), les polychloro-biphényles (PCB, isolants électriques) et d’autres xénobiotiques. AhR induit les gènes de la détoxification des hydrocarbures aromatiques tels que le cytochrome

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P450 1A1 (CYP 1A1) et les enzymes de conjugaison (glutathion S-transférase Ya, UDP-glucuronyl-transférase, etc.) [19] et un grand nombre de gènes liés à l’inflammation (interleukines, Cox2, NO synthase) [20]. Certains ligands comme les PCB augmentent le stress oxydatif et la peroxydation des lipides, ce qui pourrait être une cause des lésions cellulaires et du rôle athérogène de ces molécules. Ces composés agissent sur la vascularisation et provoquent des perturbations métaboliques conduisant à l’insulino-résistance. Un autre ligand, le TCDD inhibe la différenciation adipocytaire [21] et le transport du glucose dans le tissu adipeux [22]. Des études épidémiologiques montrent que l’exposition à la dioxine tétrachlorée (dioxine de Seveso, TCDD) induit une augmentation significative du risque de TD2. Bertazzi et al. rapportent dès 1998 un “excès” de cas de TD2 dans la population féminine de Seveso [23, 24]. Ce lien entre dioxine et TD2 a été confirmé par les études ultérieures de ce groupe [25] et par des études sur des anciens combattants du ViêtNam [26, 27] et des travailleurs exposés [28, 29]. Ces données sont à rapprocher de l’association connue entre TD2 et tabagisme, dont les agents pricipaux (hydrocarbures aromatiques) sont des ligands du AhR [30, 31]. Le RVR inhibe les effets des divers agonistes du AhR par inhibition compétitive [32]. Le RVR est le seul antidote efficace et non toxique contre ces polluants environnementaux.

Figure 1 : Structure du resvératrol

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Le resvératrol : molécule antioxydante

Le RVR est connu depuis de nombreuses décennies comme étant un antioxydant majeur. Mais il faut préciser que c’est le RVR ingéré et non métabolisé qui possède cette activité. Malheureusement, le RVR est très vite métabolisé, et les effets de cette molécule sur le syndrome métabolique proviendraient d’autres propriétés que nous décrirons ultérieurement.

Le resvératrol : un anti-inflammatoire antagoniste des voies NF- B et

AP-1.

Les cascades moléculaires liées à AhR et conduisant à une résistance à l’insuline ne sont pas identifiées mais l’implication de NF- B, facteur essentiel de l’inflammation a été suggérée [33, 34]. L’activation du AhR par les dioxines induit l’induction des cytokines de types IL-1. En inhibant NF- B par son antagonisme vis à vis du AhR, le RVR inhibe l’induction des cytokines inflammatoires. Ce composé inhibe également l’expression de la chimiokine MCP-1 [35, 36] induite par le TNF! et qui est impliquée dans l’infiltration macrophagique. Il se pourrait donc que le RVR agisse sur le DT2 via son effet au niveau de l’expression de MCP-1. Le AhR interagit physiquement avec d’autres facteurs de transcription dont PPAR" [37, 38] tandis que PPAR! régule l’expression de AhR [39, 40]. Il est important de noter que le facteur de transcription PPAR" est la cible de médicaments anti-diabétiques de la famille des thiazolidinediones et PPAR! joue un rôle dans la lipolyse. Le RVR agit aussi sur l’inflammation en inhibant les voies NF- B et AP-1 [41-45]. Il diminue l’expression de la cytokine IL-1# et s’oppose à la production de radicaux oxydants (ROS) [46, 47]. D’autres voies d’action du RVR ont été mises en avant comme la modification de la synthèse des eicosanoïdes par inhibition des cyclooxygénases (COX-1 et COX-2) responsables de la synthèse de médiateurs pro-inflammatoires [48].

Le resvératrol et les mitochondries

Dans le muscle squelettique, une diminution de la synthèse d’ATP et de l’expression des gènes contrôlant l’activité mitochondriale est en partie, au moins, associée à une insulinorésistance. Le facteur PGC-1!$qui est un PPAR"$et le facteur SIRT1 qui est associé à PGC-1 sont fonctionnellement liés l’un à l’autre et à l’utilisation du glucose. En effet, une diminution de l’activité de PGC-1 est associée à l’apparition du diabète et SIRT contrôle le métabolisme glucidique hépatique [49].Le RVR induit l’expression du gène codant pour PGC-1! via l’activation du facteur SIRT1 [49]. Par ailleurs, le RVR augmente la taille et le nombre de mitochondries dans divers tissus. Dans les muscles, en réponse au RVR, les mitochondries présentent une augmentation du pouvoir oxydant et une activité métabolique accrue associée à une augmentation de l’activité de la citrate synthétase.

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Figure 2 : Étude pharmacocinétique réalisée chez 15 sujets sains en milieu hospitalier. 40 mg de RVR sont administrés soit sous forme sèche (classique) soit sous forme liquide (formule du laboratoire). Le taux de RVR

sanguin est évalué au cours du temps. Le resvératrol sous forme liquide est toujours dans une concentration thérapeutique active 5 heures après son administration.

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LES DONNÉES DE NOTRE LABORATOIRE

Notre laboratoire s’est impliqué dans l’étude de l’effet d’un traitement au RVR sur le TD2. Le RVR, dès son administration chez l’homme, est une molécule qui est très vite éliminée de l’organisme suite à un cycle de métabolisation entéro-hépatique très fort impliquant les enzymes UGT1A1 et 1A9. Cela se traduit par l’apparition de métabolites sulfatés et de glucuronoconjugués éliminés par les urines et les fèces.Avant de valider l’effet du RVR sur le TD2, nous avons mis au point une forme galénique permettant au RVR d’être hautement biodisponible [50]. Après validation de cette galénique par des études de pharmacocinétique (figure 2), nous avons nourri des animaux par un régime diabétogène riche en graisse supplémenté ou non en RVR.

L’administration de RVR pendant 5 semaines réduit l’intolérance au glucose. Nous avons mis en évidence les mécanismes moléculaires impliqués dans ce processus. Nous avons démontré que le RVR induisait, par augmentation de la transcription, l’expression et ensuite la sécrétion intestinale de Glucagon-Like Peptide-1 (GLP-1) [51]. Cette hormone entre par la veine porte et peut alors activer la sécrétion d’insuline. Ce mécanisme a été validé par l’utilisation d’animaux génétiquement modifiés et présentant un récepteur

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Figure 3 : Effet de RVR sur la flore intestinale. NC : contrôle ; INFL : régime inflammatogène ; INFL + RVR : régime inflammatogène + resvératrol. Le RVR fait disparaître des bactéries qui ont émergé suite au régime

inflammatogène et fait apparaître des bactéries qui ont été tuées par ce régime.

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non fonctionnel au GLP-1. En effet, sur ces animaux, le RVR n’agit plus sur la tolérance au glucose car le taux d’insuline n’est plus modulé. Parallèlement à cela, nous avons démontré que le RVR diminue l’inflammation induite par le régime diabétogène riche en graisse. En effet, l’expression des cytokines inflammatoires de type IL-1 et IL-6 est diminuée par le RVR et les cytokines anti-inflammatoires de type IL-10 et TGF sont augmentées par la même molécule en particulier au niveau hépatique et intestinal [51]. Le RVR étant une molécule antifongique qui possède quelques propriétés antimicrobiennes, nous avons évalué l’effet du RVR sur la flore intestinale. Nous démontrons pour la première fois que le RVR est un agent prébiotique qui diminue le nombre de bactéries inflammatogènes Gram - et qui induit la prolifération des bactéries Gram + (figure 3) [51].

Enfin, nous avons testé l’effet d’une association du RVR à la sitagliptine, un médicament antidiabétique inhibiteur de la dipeptidyl peptidase-4 (DPP-4). Ce choix fut motivé par le fait que le RVR induit la sécrétion de GLP-1 et que la sitagliptine inhibe la dégradation du GLP-1. Nos résultats indiquent finalement que le resvératrol potentialise l’effet de la sitagliptine [51].

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CONCLUSION

Le RVR sous une forme biodisponible par voie orale, en tant qu’«alicament», semble un candidat prometteur dans le domaine du traitement du TD2 et de l’obésité. Toutes les données cumulées sur cette molécule confirment que cette molécule peut être utilisée dans la prévention ou l’optimisation d’une thérapeutique antidiabétique. L’avantage du RVR est d’être un complément alimentaire naturel et non toxique, utilisable per os et associable à un médicament anti-diabétique. Son activité sur le TD2 est médié par ses divers rôles anti-inflammatoire, prébiotique et inducteur de la sécrétion du GLP-1 et de l’insuline.

Dr Eric SERÉE

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