Sphingolipids, rafts and multidrug resistance Hinrichs, Joann Wilhelm Jakob

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1 Sphingolipids, rafts and multidrug resistance Hinrichs, Joann Wilhelm Jakob IMPORTANT NOTE: You are advised to consult the publisher's version (publisher's PDF) if you wish to cite from it. Please check the document version below. Document Version Publisher's PDF, also known as Version of record Publication date: 2004 Link to publication in University of Groningen/UMCG research database Citation for published version (APA): Hinrichs, J. W. J. (2004). Sphingolipids, rafts and multidrug resistance s.n. Copyright Other than for strictly personal use, it is not permitted to download or to forward/distribute the text or part of it without the consent of the author(s) and/or copyright holder(s), unless the work is under an open content license (like Creative Commons). Take-down policy If you believe that this document breaches copyright please contact us providing details, and we will remove access to the work immediately and investigate your claim. Downloaded from the University of Groningen/UMCG research database (Pure): For technical reasons the number of authors shown on this cover page is limited to 10 maximum. Download date:

2 Chapter 8 Nederlandse samenvatting John W. J. Hinrichs Groningen University Institute for Drug Exploration, Department of Membrane Cell Biology University of Groningen, The Netherlands 141

3 Ons lichaam is opgebouwd uit miljarden cellen die ontstaan zijn uit één bevruchte eicel door celdeling: uit één cel ontstaan twee nieuwe cellen, die zich op hun beurt ook weer delen, enzovoort. Behalve voor de groei is celdeling ook noodzakelijk om beschadigde en oude cellen te vervangen. In normale omstandigheden wordt de celdeling streng gereguleerd door specifieke genen die onderdeel zijn van het DNA dat zich in elke lichaamscel bevindt. Deze zogenaamde proto-oncogenen en tumor-suppressor genen maken stoffen aan die de celdeling stimuleren, respectievelijk remmen. Bij kankercellen is het DNA van deze regelgenen beschadigd, waardoor de cel te veel of juist te weinig van deze stoffen aanmaakt. Het resultaat is in beide gevallen hetzelfde: ongeremde celgroei ofwel kanker. In geval van chemotherapie worden deze ongeremd delende cellen behandeld met speciale medicijnen, cytostatica genoemd. Deze cytostatica remmen de celdeling en omdat kankercellen zich zeer snel delen worden deze hierdoor het meest getroffen. Een belangrijk probleem bij chemotherapie is echter dat kankercellen na verloop van tijd ongevoelig kunnen worden voor deze medicijnen. De tumor wordt resistent voor de therapie en kan gewoon doorgroeien. In veel gevallen zijn de kankercellen niet alleen ongevoelig voor het oorspronkelijk gebruikte medicijn, maar ook voor andere structureel niet gerelateerde medicijnen. Deze zogeheten multi-drug resistentie (MDR) maakt de verdere behandeling van de tumor erg lastig. Er zijn diverse redenen voor MDR, maar de voornaamste reden is de opregulatie van zogenaamde drug-efflux pompen in kankercellen. Dit zijn grote eiwitten die zich bevinden op het grensvlak tussen binnen en buiten de cel, het zogenaamde plasmamembraan. Deze drug-efflux pompen kunnen diverse moleculen, waaronder cytostatica, effectief uit de cel transporteren. Als onderdeel van het plasmamembraan hebben deze eiwitten intensief contact met de bouwstenen van dit membraan, namelijk de lipiden. Lipiden bestaan er in allerlei soorten en maten, maar ze hebben allemaal een hydrofiele (waterminnende) kop en een hydrofobe 142

4 (waterafstotende) staart. De plasmamembraan bestaat eigenlijk uit twee lagen geordende lipiden, die gespiegeld op elkaar liggen, met de waterafstotende staarten naar elkaar toegekeerd en de waterminnende koppen naar de omgeving gericht. Dat resulteert in een hydrofobe tussenlaag en een hydrofiele buitenkant. Het is al bekend dat drug-efflux pompen beter of slechter werken afhankelijk van hun lipiden omgeving. De functie van de pompen wordt dus beïnvloed door de soort lipiden waarmee ze in contact staan. Behalve opgereguleerde drug-efflux pompen, vertonen MDR kankercellen vaak een specifieke toename van een bepaald soort lipiden. Deze zogenaamde sfingolipiden zijn een belangrijk onderdeel van specifieke domeinen in de plasmamembraan. Deze domeinen worden ook wel rafts genoemd. Het plasmamembraan is geen statisch geheel, maar is dynamisch. De verschillende eiwitten en lipiden kunnen ten opzichte van elkaar bewegen binnen de lipidenlagen. Zo kunnen clusters ontstaan van specifieke lipiden en eiwitten die zich fysisch en functioneel onderscheiden van hun omgeving in het vlak van de membraan. Deze rafts zijn te isoleren uit de membranen op basis van onoplosbaarheid in detergentia. Detergentia zijn moleculen die lipidenmembranen kunnen oplossen, wat betekent dat ze de onderlinge bindingen tussen de lipiden kunnen verbreken. Rafts onderscheiden zich van de rest van het plasmamembraan, doordat ze sfingolipiden bevatten die onderling zeer sterke verbindingen vormen, die door de detergentia niet kunnen worden verbroken. Behalve een verrijking van sfingolipiden in deze membraandomeinen zijn ook drug-efflux pompen geconcentreerd in deze domeinen. In dit proefschrift wordt beschreven dat drug-efflux pompen en sfingolipiden gelijktijdig worden op-gereguleerd tijdens de onwikkeling van MDR in kankercellen. Bovendien speelt deze gezamenlijke op-regulatie zich hoofdzakelijk af in membraandomeinen. De drug-efflux pompen blijken zich te bevinden in een specifiek raft-type. Dit raft-type wijkt af van het 143

5 klassieke type raft (lees: het eerst ontdekte type) op basis van de lipidensamenstelling. In tegenstelling tot het klassieke raft-type bevat het een relatief grote hoeveelheid van die lipiden, die noodzakelijk zijn voor de werking van de drug-efflux pompen. Het merendeel van de sfingolipiden daarentegen lijkt zich te concentreren in de klassieke rafts. Het blijkt echter ook dat beide raft-typen grotendeels overlappen. Op basis van de verkregen resultaten en gegevens uit de literatuur postuleren we een model dat drug-efflux pompen én sfingolipiden functioneel combineert in één samengestelde raft. Naar verwachting zullen cytostatica zich, op basis van hun chemische eigenschappen, concentreren in de sfingolipidenrijke kern van de raft. De drug-efflux pompen, gelegen in een voor hun functie optimale lipidenomgeving, vormen hieromheen een soort cordon, dat weglekkende cytostatica effectief uit de cel kan verwijderen. 144

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