Mechanistic aspects and structural organization of a secondary sugar transporter
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- Bernard van der Berg
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1 Mechanistic aspects and structural organization of a secondary sugar transporter
2 ii Cover: entries and exits (Oosterpoort, Groningen) The study described in this thesis was carrier out in the Departments of Microbiology and Biochemistry of the Groningen Biomolecular Sciences and Biotechnology Institute (GBB) of the University of Groningen. The work was financially supported by E.U. grants BIO4-CT , and and a Jonge Chemici grant from the Netherlands Organization for Scientific Research (NWO) under the auspices of the Council for Chemical Sciences. Printed by: PrintPartners Ipskamp, Enschede ISBN:
3 iii RIJKSUNIVERSITEIT GRONINGEN Mechanistic aspects and structural organization of a secondary sugar transporter Proefschrift ter verkrijging van het doctoraat in de Wiskunde en Natuurwetenschappen aan de Rijksuniversiteit Groningen op gezag van de Rector Magnificus, dr. D.F.J. Bosscher, in het openbaar te verdedigen op vrijdag 7 september 2001 om uur door Liesbeth Marian Veenhoff geboren op 23 november 1972 te Groningen
4 iv Promotor: Prof. Dr. B. Poolman Beoordelingscommissie: Prof. Dr. A.J.M. Driessen Prof. Dr. W.N. Konings Prof. Dr. G.T. Robillard ISBN:
5 v Bedankt! Het proefschrift is af: een mooi moment om een aantal mensen te bedanken die elk op eigen wijze betrokken zijn geweest. Allereerst wil ik mijn promotor Bert Poolman bedanken. Je enthousiasme voor het onderzoek, je mammoetgeheugen, je vele ideeën, je kritische blik op wetenschappelijke teksten en je nieuwsgierigheid heb ik altijd zeer gewaardeerd. Met de vrijheid die je me liet en het vertrouwen dat je had in mijn werk, is mijn promotieonderzoek geworden wat het is. Veel dank hiervoor! Wil Konings wil ik bedanken voor de tijd dat ik werkte in zijn befaamde Koninkrijk, waar hij met overtuiging heer en meester was van een indrukwekkende onderzoeksgroep. Aan LacS werkten we op de hoogtijdagen met 6 mensen en het leek soms of het eiwit domweg geen zin had om zijn geheimen prijs te geven, maar we zijn toch een heel eind gekomen. Jan en Robert, dankzij jullie werk aan LacS ontstonden vele nieuwe mogelijkheden waarvan ik dankbaar gebruik heb gemaakt. Brutus (Frits ), Mark en Eric, bedankt voor de vele proeven en veel succes! Ria, ik kijk met plezier terug op de periode dat we samen met Esther werkten aan hoofdstuk 5 in dit boekje, bedankt! Marga, je bent in vele opzichten een supervoorbeeld voor me! Wetenschap is een mooi vak, maar wat nou precies maakt dat een lab zo n bijzondere werkplaats is? Misschien is het omdat er naast het doen van experimenten veel ruimte is voor grappen en onzinverhalen bij het doen van alle domme klussen, of omdat de moeizame periodes vragen om een borrel en saamhorigheid. De vele promoties die groots gevierd moesten worden, en de bonte stoet van mensen die kwamen en gingen hebben zeker bijgedragen. Ik zal nog vaak met weemoed terugdenken aan de vele cabarets, etentjes, promotiefeesten, de (wilde) borrels en barbecues, de wandel- en zeiltochtjes, de lauwersloop dagen, en alle grappen op het lab. Ik vond het bere-gezellig met jullie, erg bedankt! Het is alweer wat langer geleden maar ik ben jullie nog lang niet vergeten, de mooie oude gekken van het lab: Jack, Marleen, Erik, Jeroen, André en Janine, Ted, Theo en Mike. Jessica en Bastiaan, jullie hebben de boel goed draaiende gehouden in Haren. Een paar mensen die de laatste periode voor veel steun en gezelligheid hebben gezorgd wil ik met name noemen, Marga (een held, zoals ik al schreef), Frank (onze poëet en trouwe drinkerbroeder), De Dirkjes en Janny (altijd scherp en geestig), Sonja (Lunteren is je lot, meid!), Tiemen (je hebt je kranig geweerd!) en Gert (onze enige echte wetenschapper en liedtekstschrijver), bedankt! Hoop dat we elkaar niet helemaal uit het oog verliezen!
6 vi Esther, deze eerste paar jaar in de wetenschap gaf voor mij een hoop te denken en te bespreken, en ik heb het als een enorme steun en geruststelling ervaren te merken dat we het meestal roerend eens waren. Met z'n tweeën heb je toch net iets meer gelijk, en menig deur hebben we getweeën toch maar mooi open gekregen! Ik ben er trots op dat we twee hoofdstukken samen hebben geschreven, en heb er met veel plezier aan gewerkt. Waaaaaaaahnzinnig bedankt, alle helden in mijn leven! Zonder jullie zou er geen biet aan zijn. Ik heb veel van jullie geleerd, en jullie bijdrage aan dit proefschrift is dan ook groots. Ik ben stikgelukkig met jullie allemaal, lieve Monika en Dick, Bert en Anke, Anneke en Floris, Oma, Maaike en Morris, Femke en Rick, Margreet en Gjalt, Jildou, Hans en Wietze, Theo en Linda, Caro, Silke en Wil, Rahel, Susanne en Richard, Mags en Gigi, Barth, Esther, Saskia, Corlien, Anke, Willem, Ilja, Margriet, Reinier en, sinds kort, Tom. Maaike en Femke, onze vriendschap is mij heel dierbaar, ook nu we nog maar weinig hutten bouwen in het bos! Lieve Monika, Bert en Anneke: dat in mijn herinneringen aan vroeger de tuin altijd bloeide, de piano pingelde, de schommel nooit stil stond, de bergen witbesneeuwd waren en de koekjes net uit de oven kwamen, is een mooi bezit. Bedankt voor het vele meedenken met alles! JJ, mijn lief,... een kus voor jou! Liesbeth
7 vii contents page Chapter 1: Chapter 2: Chapter 3: Chapter 4: Chapter 5: Chapter 6: Chapter 7: Chapter 8: Dankwoord General introduction and scope of this thesis Substrate recognition at the cytoplasmic and extracellular binding site of the lactose transport protein Proximity of putative α-helices II, IV, VII, X and XI in the translocation pathway of the lactose transport protein The lactose transport protein is a cooperative dimer with two sugar translocation pathways Oligomeric state of membrane transport proteins analyzed with blue native electrophoresis and analytical ultracentrifugation Quaternary structure and function of secondary transport proteins Summary and concluding remarks Nederlandse samenvatting (voor niet-biologen) References Abbreviations List of publications
8 viii
9 Chapter 1 General Introduction and Scope of this Thesis Contents: Page Why study membrane transport by LacS 10 Classes of membrane transport proteins 10 The GPH-family of Major Facilitators 12 The Major Facilitator Superfamily The Galactoside-Pentoside-Hexuronide(GPH):cation symporter family The lactose transport protein of Streptococcus thermophilus 13 Biological function Transport mechanism Regulation of transport activity Structure Experimental tools Outline of the thesis 17
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