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Major Histocompatibility Complex Genes in the Common Carp, Cyprinus Carpio L

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Major Histocompatibility Complex Genes in the Common Carp, Cyprinus Carpio L Major histocompatibility complex genes in the common carp, Cyprinus carpio L. Promotor: dr. W.B. van Muiswinkel hoogleraar in de zoölogische celbiologie Co-promotoren: dr. R.J.M. Stet universitair docent in de zoölogische celbiologie/immunogenetica dr. E. Egberts universitair hoofddocent in de zoölogische celbiologie/immunologie M N0®2©\ ,200« Major histocompatibility complex genes in the common carp, Cyprinus carpio L. Saskia H.M. van Erp Proefschrift ter verkrijging van de graad van doctor in de landbouw- en milieuwetenschappen, op gezag van de rector magnificus, dr. C.M. Karssen, in het openbaar te verdedigen op woensdag 29 mei 1996 des namiddags te half twee in de aula van de Landbouwuniversiteit te Wageningen AISV'U CIP-GEGEVENS KONINKLIJKE BIBLIOTHEEK, DEN HAAG Erp, Saskia H.M. van Major histocompatibility complex genes in the common carp, Cyprinuscarpio L. / Saskia H.M. van Erp. - [S.l. : s.n.]. -111 . Thesis Landbouwuniversiteit Wageningen. - With réf. - With summary in Dutch. ISBN: 90-5485-509-6 Subject headings: carp / major histocompatibility complex / comparative immunology. Abstract VanErp , S.H.M. 1996. Major histocompatibility complex genes in the common carp, Cyprinus carpio L. Doctoral thesis, Dept. Experimental Animal Morphology and Cell Biology, Wageningen Agricultural University, P.O. Box338 , 6700A HWageningen , TheNetherlands . This thesis describes astud y of themajo r histocompatibility complex (Mhc) genes of thecommo n carp (Cyprinus carpio L.). Themolecule s encoded byMhc genes play anessentia l role inth e specific immune response, by presenting antigens to T lymphocytes. Knowledge of theMhc of carp, therefore, contributes to our understanding of the immune responsemechanism s inthi s species. In addition, itma y give important insights inth ephylogeneti c development ofthes e genes. The common carp was found to contain several distinct lineages ofMhc class Igenes , denoted as Cyca-U, Cyca-Z, Cyca-TC16an d Cyca-CA. The Cyca-U sequences probably represent classical Mhc class Igenes , of which most likely only asingl elocu s is expressed ineac h individual. Cyca-Z genes are present inmultipl epolymorphi c copies inth e genome, but it isno t clear whether these genes are expressed. The sequence of Cyca-TC16 ismos t similar to the classI genes of the coelacanth, afish whic h isthough tt ob e arepresentativ eo f the evolutionarylineag e leading to the tetrapods. It is, however, not clear whether Cyca-TC16 isexpressed . In addition, the sequence encoding carp ß2-microglobulinwa s isolated. Althoughtw oB2m genes weredetecte d in each individual, apparently only one of these isexpressed . In contrast, at least four class IIB genes may be expressed in a singleanimal . These genes are linked intw opairs , which, however, segregate independently. In addition, twoexpresse d class IIA sequences were identified, most likely derived from two separate loci. Bothth e class IIA andB genes are likely to encode bona fide class II chains, the components of the cell-surface class II a-ß heterodimer. Althoughcar p thusposses s a complete set of class I and class II genes, it is not yet clear whether thesegene s reside in asingl egeneti cMhc region. Cover: Drie werelden. ® 1996 M.C. Escher / Cordon Art - Baarn - Holland. Alle rechten voorbehouden. *ƒ)» ^JÖÖW^S? Stellingen 1. De aanwezigheid van zowel Mhc klasse I als klasse II genen in de karper wijst erop dat al bij vissen de strategische tweedeling aanwezig is tussen immuunresponsen tegen intra- en extracellulaire antigenen. Waarschijnlijk zijn er dan ook CD4- en CD8-achtige T-cel co-receptoren aanwezig die hierin een sturende rol spelen. ditproefschrift 2. Naamgeving aan Mhc genen is gelijk de kip en het ei: een nomenclatuur die de relaties tussen genen helder enjuis t weerspiegelt is van groot belang voor het succes van het onderzoek, maar zij kan slechts juist zijn wanneer ditzelfde onderzoek de relaties volledig heeft opgehelderd, ditproefschrift 3. In termen van zijn Mhc is hetjuiste r de karper evolutionair gezien een vroegere vertebraat te noemen dan een lagere, ditproefschrift 4. Twijfel is een eerbetoon aan de waarheid - Ernest Renan - 5. In evolutionair opzicht is het ongunstig als carrièrevrouwen kinderloos blijven. - W.A.M, van Erp - 6. Dat karpers een hoge aaibaarheidsfactor hebben kan experimenteel worden aangetoond in het hoofdgebouw van de Landbouwuniversiteit Wageningen. 7. AIO's zijn alsjong e hondjes: Hoe vaak ze ook kwispelend al experimenterend tegen een boom aan lopen, na de ergste duizeligheid kwispelen ze alweer. 8. Het is een schrijnende tegenstelling dat voortschrijdende verlichting door de wetenschap juist geloof in het occulte lijkt te stimuleren. 9. Bezuinigingen die tot gevolg hebben dat wetenschappers eigenhandig boekenplanken voor hun kamer moeten kopen en ophangen, leveren waarschijnlijk alleen de Gamma geld op. 10. Hoogstwaarschijnlijk is de volksgezondheid meer gebaat bij een radicale vernietiging van sigaretten dan van Brits rundvee. Uit het uitblijven van eerstgenoemde maatregel kan dan ook worden opgemaakt dat een gezonde economie al snel belangrijker is dan gezond volk. Stellingen behorend bij hetproefschrift "Majorhistocompatibility complex genes inthe common carp (Cyprinus carpio L.)" van Saskia H.M. van Erp, Wageningen, 29 mei 1996. Voorwoord Mijn proeve van bekwaamheid. Die houdt U vast. U snapt hoe bekwaam ik me voel met dit boekje. Maar dit werkje betekent veel meer dandi e proeve alleen. Het staat voor 5jaa r "nog even naar het lab gaan", voor 5jaa r adrenaline, vreugde en veel ontzetting. Zo gaat dat in de wetenschap. En gelukkig was daar altijd weer het "En? Is het gelukt?" van een van de menseno mm eheen . Zonder hunaanwezighei d zoud eMajo r Hysterical Crisishee lwa thoge r tij hebben gevierd. Om te beginnen natuurlijk René, een rots in de branding, een baken op zee. Met bijbehorend aanvaringsgevaar natuurlijk. Maar samen hebben we het gered, René. Dankjewel. Ennatuurlij k ookEgbert ,Brian ,Pedro ,Trud ie nMarie-José .He twa sgoe dwerke n engoe d toeven injulli egezelschap . Wim, dankj e wel voorj e blinde vertrouwen inhe t succes van mijn werk. De AIO's van EDC (inclusief doctor Geert) had ik voor geen goud willen missen. Veel eten, veel wijn, veel idealen. De wereld verklaard, de ziel gesterkt. Mooie avonden. Ook dank aan de studenten die slechts achteraf genoemd in dit boekje voorkomen. Erike nJoris ,julli eware ngezellig emaatje s ophe tlab .I khe bminsten szovee lva njulli egeleer d als jullie (hopelijk) van mij. Robert, Jeroen en Bart, hoewel slechts vanaf de zijlijn heb ik van jullie werk dankbaar enkele belangrijke graantjes meegepikt. Jan (Osse), jouw belangstellende bezoekjes aan het eind van een werkdag heb ik erg gewaardeerd. En verder natuurlijk de rest vanEDC , dankjulli e wel voorjulli e interesse, koffiepraat, valentijnsrozen, gezelschap, enz. enz. Minstensd ehelf tva nmij nleve nal sAI Oha di khe tgenoege nt evertoeve ntusse nd e Veefokkers op het MHC lab. Richard, je bent van goud. Je was onmisbaar voor me, inzowe l praktische als persoonlijke zin. Jan, Martien, Tony (ik misj e confronterende grappen nog steeds, "Ik dachtda t ikee nkamee lhoorde....") , Carolien, Beja, Rosilde, Tineke, Esther, enall e anderen die in de jaren kwamen en gingen, ik heb genoten injulli e midden. Peter, jouw opgewektheid is legendarisch. Dankjewel voor alle zonnige hulp aan de copiëermachine. Wim Valen, dankj e wel voor alle plaatjes die je in dejare n voor me hebt geschoten. Mijn paranimfen, Gertjan enRichard , dankjulli evoo r alle steune ngoed e zorgen. A few words of special thanks to Felipe Figueroa and JanKlein , for their hospitality, support and volleyball-training during my stay in Tübingen. Papae nmama , Florise nJacintha , dankjulli evoo rd eliefdevoll e basise nhe t onvoorwaardelijk vertrouwen date r altijd voorm e is. Paul,j e weetondertusse n evenveelva nPC R enallelisch e lijnen als ik, ik hoop dat je nog lang mijn onderwijzerskwaliteiten zult waarderen. Dank je voor alle geduld en liefde. Enda nzij ndaa rno gd ehoofdrolspelers . Stilo pd eachtergrond ,lijden dvoorwer pe nonderwer p tegelijk. De karpers. Ze hebben geen idee wat voor geheimen ze hebben prijsgegeven. Zelfs niet die ene op de voorkant, met zijn neus er bovenop. Dank je wel visjes! Contents Chapter 1 General introduction Chapter 2 Polymorphism and estimation of the number ofMhcCyca class I and class II genes in laboratory strains of the common carp (Cyprinuscarpio L.) 17 Chapter 3 Characterization of ß2-microglobulin transcripts from two teleost species 39 Chapter 4 Identification and characterization of a new major histocompatibility complex class I gene in carp {Cyprinuscarpio L.) 59 Chapter 5 Evidence for multiple distinct major histocompatibility complex class I lineages in teleostean fish 89 Chapter 6 Characterization of major histocompatibility complex class IIA and B genes in a gynogenetic carp clone 107 Chapter 7 General discussion 131 Summary 147 Samenvatting 153 Curriculum vitae 159 Chapter 1 General introduction Allwarm -an dcold-bloode dvertebrate sposses sa mor eo rles sintegrate dimmun esyste m by which the organism defends itself against unwanted invaders like parasites, bacteria, and viruses, butals oagains tdanger s from within, liketumo rcells . Harmful attacksb yth eimmun e system on self antigenshav e to beavoided , however, and amechanis m should therefore exist by which the immune system candiscriminat e between self andnon-self . This discrimination is performed by T lymphocytes which are selected early in their development for tolerance to self, but reactivity to non-self. Antigen recognition by T cells is performed through their T-cell receptors which, however, can only recognize antigens when these are presented as peptides by Mhc molecules, the transmembrane glycoproteins encoded by the genes of the major histocompatibility complex {Mhc). In mammals, there are two major classes of Mhc molecules, class I and class II, which are distinguished by their protein structure, function and tissue distribution. Mhc class I molecules are heterodimers of a heavy class I a chain (Mr 45,000), complexed with a light chain, ß2-microglobulin (Mr 12,000).
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