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Phd the Fountain of Youth Final 3 Juni Ghent University Faculty of Sciences Academic year 2010-2011 The quest for the fountain of youth: Developing flatworm models for ageing and rejuvenation research Stijn Mouton Promotors: Prof. Dr. Bart Braeckman (UGent), Dr. Wouter Houthoofd (GlaxoSmithKline Biologicals), Dr. Maxime Willems (UGent) Thesis submitted to obtain the degree of doctor in Sciences (Biology) Proefschrift voorgelegd tot het bekomen van de graad van doctor in de Wetenschappen (Biologie) 1 2 Reading committee Prof. Dr. Lukas Schärer (University of Basel, Switzerland) Prof. Dr. Tom Artois (Hasselt University, Belgium) Prof. Dr. Wim Bert (Ghent University, Belgium) Examination committee Prof. Dr. Dominique Adriaens (chairman, Ghent University, Belgium) Prof. Dr. Lukas Schärer (University of Basel, Switzerland) Prof. Dr. Tom Artois (Hasselt University, Belgium) Prof. Dr. Wim Bert (Ghent University, Belgium) Prof. Dr. Bart Braeckman (Promotor, Ghent University, Belgium) Dr. Wouter Houthoofd (Promotor, GlaxoSmithKline Biologicals, Belgium) Dr. Maxime Willems (Promoter, Ghent University, Belgium) 3 4 Dankwoord – Acknowledgements Een doctoraat schrijf je niet alleen, maar met de hulp en steun van heel veel mensen. Daarom een grote merci aan iedereen die op een of andere manier bijgedragen heeft aan dit boek. In de eerste plaats mijn promotors en lableider. Prof. dr. Bart Braeckman, bedankt voor het ter beschikking stellen van je lab en de nodige apparatuur voor het uitvoeren van de metabolische experimenten, het verwerken van alle worm-foto’s, het zo vaak grondig verbeteren van mijn Engels en de hulp bij het interpreteren van de TAM data. Het niveau van het TAM artikel is dankzij jou enorm verbeterd (zeker in vergelijking met mijn eerste versie) en ik heb ongelooflijk veel bijgeleerd door samen aan dit manuscript te werken. Dr. Wouter Houthoofd, vooral merci om altijd klaar te staan, te motiveren en te antwoorden op allerhande vragen. Al toen ik 18 was, kwam ik bij jou uitleg vragen over de richting Biologie, en het is deels door jouw enthousiasme toen dat ik uiteindelijk hiervoor gekozen heb. Ook tijdens mijn doctoraat stond je altijd klaar om te helpen waar nodig (planningen opstellen is er zeker een van) en je bent steeds een belangrijke bron van motivatie geweest. Merci ook voor alle leuke momenten naast het werk, en je enorme gastvrijheid als ik in Leuven bleef logeren. Prof. dr. Wim Bert, je hebt vooral een grote rol gespeeld bij de schrijffase van dit boek. Merci voor je steun bij het worstelen met de statistiek, de hulp met de figuren, het ongelooflijk snel antwoorden op 5 mijn stress-mails, het pdf’en van deze versie en de organisatie van het printen en drukken van dit boek. En natuurlijk Dr. Maxime Willems, ik weet niet waar te beginnen met je te bedanken. Je bent mijn begeleider geweest vanaf de bachelorproef tot het einde van mijn doctoraat en zonder jouw hulp was dit boek er zeker nooit gekomen. Zonder jou was er zelfs niet eens platworm-onderzoek in Gent. Je invloed op mijn doctoraat is dan ook gigantisch. Merci voor de talloze wetenschappelijke discussies, de hulp bij het maken van figuren, het samen schrijven van artikels, het bedenken van experimenten, het motiveren waar nodig maar ook het relativeren waar nodig, en zo veel meer. Ik hoop echt dat we in de toekomst nog samen kunnen werken aan het ontwikkelen van Macrostomum als model organisme In de rij van de begeleiders zou ik ook dr. Lukas Schärer willen bedanken. Lukas, thanks for all the scientific discussions, organising the yearly Macrostomum meetings, your (and Dita’s) hospitality when I was staying at your place, and the ungoing help with the “Back to the future-article”. You had a big influence on the research described in this book. Verder zijn er natuurlijk de mensen van het lab. Een enorme merci voor Marjolein. Ook jij was essentieel in het bekomen van de resultaten van dit doctoraat. Merci voor het maken van medium en het gieten van platen (ik vraag me af hoeveel je er gegoten zou hebben in totaal), het maken van alle producten, de hulp bij de respiratiemetingen en het op punt stellen van de maceratie en 6 flow cytometry. Als jij ergens enthousiast over was, ging alles plots dubbel zo snel vooruit. Ik heb je al gemist in het lab in Utrecht. Freija, merci voor de hulp bij het wetenschappelijke werk, maar ook voor de goede sfeer van ons lab-eiland en zeker merci voor de amazon-bestellingen. Tijd dat we nog eens cd’s uitwisselen. Mieke, bedankt voor het nalezen van teksten, de vele lay-out tips voor presentaties en poster en zeker ook voor het maken van de uitnodiging voor de verdediging. Hanne en Pam, merci voor alle leuke momenten in het lab. Jeroen en Ineke, jullie waren de perfecte bachelorstudenten. Nog eens Jeroen , merci voor het vele werk dat je in je thesis gestoken hebt. Je enthousiasme maakte het begeleiden van een thesis een van de leukste dingen tijdens mijn doctoraat. Myriam, merci voor de hulp met de TEM. Nog een aantal mensen die moeten bedankt worden zijn: Leen Leus, voor de hulp bij de flow cytometry; Greet en Frederik, voor de hulp met de statistiek, en Kristina voor het maken van de cover van dit boek. Verder wil ik iedereen van het derde verdiep en het ageing lab bedanken voor de supergoede sfeer. Ik zal de gezellige middagen in de plantentuin, de filmavonden in de Ledeganck, de quiz, de vele etentjes, enz. nog lang herinneren. Dankzij die goede sfeer was het altijd leuk om te komen werken en leek het doctoraat vaak meer een hobby dan werk. 7 Speciaal wil ik ook Heleen en Patricia bedanken. Jullie stonden altijd klaar voor een babbel en doorheen de jaren zijn jullie goede vrienden geworden. Ik hoop dat we elkaar nog vaak zullen zien in de toekomst. Ook naast het werk wil ik een aantal mensen bedanken. Valentijn, merci voor de goede tijd in de Tweebruggenstraat (24 periodes, kerstdinners, …), het zo vaak naar de winkel gaan als ik weer eens te laat van het lab toekwam en het regelen van alle papierwerk. De Planets: Filip, Jurgen, Guillaume en Liesbeth, merci voor de repetities. Het was een ideale uitlaatklep om het werk even te vergeten. Het ligt nu wat stil, maar hopelijk lukt het in de toekomst om terug te jammen. Ruben, Kobe, Celine en Wouter voor het opstarten van de weekend- traditie, en de gezellige zaterdagavonden. Ook daar hebben we vaak over het werk gepraat en hebben jullie altijd voor de nodige steun gezord. De rest van mijn maten voor de nodige ontspanning na het werk. Een heel speciale merci ook voor mijn familie: ma, pa, Mieke en Ridvan. Jullie zijn altijd heel geïnteresseerd geweest en zijn mijn blijven steunen en motiveren. Merci. Ten slotte nog een merci aan iedereen die ik momenteel aan het vergeten ben… 8 Table of Contents 1 General Introduction .................................................................................. 13 1.1 Ageing: a complex process ..................................................................... 14 1.2 Decreased stem cell functionality and homeostasis as causes of ageing 15 1.2 Decreased stem cell functionality and homeostasis as causes of ageing 16 1.3 The need for new model systems ............................................................ 17 1.4 Flatworms as potential new ageing models ............................................ 18 1.5 Current knowledge of flatworm ageing .................................................. 25 1.6 Flatworm rejuvenation: reversing the ageing process ............................. 28 1.7 Choosing flatworm species as potential models ..................................... 29 1.8 Aims of the study .................................................................................... 33 1.9 Outline .................................................................................................... 34 1.10 Reference list of the introduction .......................................................... 36 2 The free-living flatworm Macrostomum lignano: a new model organism for ageing research* ...................................................................................... 45 Abstract......................................................................................................... 46 Introduction .................................................................................................. 46 Macrostomum lignano: description .............................................................. 48 Life history ................................................................................................... 50 Chronological and replicative ageing ........................................................... 51 Experimental toolbox ................................................................................... 52 Lifespan extension and rejuvenation ............................................................ 55 Conclusion .................................................................................................... 60 Acknowledgements ...................................................................................... 60 Tables and Figures ........................................................................................ 61 Reference list ................................................................................................ 68 3 Demographic analysis reveals gradual senescence in the flatworm Macrostomum lignano* ................................................................................ 73 Abstract......................................................................................................... 74 9 Findings .......................................................................................................
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