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A PONTY (Cyprinus Carpio L.) PIKKELYMINTÁZAT ÖRÖKLŐDÉSI MODELLJÉNEK REVÍZIÓJA

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A PONTY (Cyprinus Carpio L.) PIKKELYMINTÁZAT ÖRÖKLŐDÉSI MODELLJÉNEK REVÍZIÓJA PANNON EGYETEM GEORGIKON KAR KESZTHELY Állattudományi és Állattenyésztéstani Tanszék DOKTORI (Ph.D.) ÉRTEKEZÉS Állat- és Agrárkörnyezet-tudományi Doktori Iskola Iskolavezető: Dr. Anda Angéla, D.Sc. (Jogelőd: Állattenyésztési Tudományok Doktori Iskola Iskolavezető: Dr. Szabó Ferenc D.Sc.) A PONTY (Cyprinus carpio L.) PIKKELYMINTÁZAT ÖRÖKLŐDÉSI MODELLJÉNEK REVÍZIÓJA Témavezetők: Dr. habil. Bercsényi Miklós egyetemi tanár Dr. Nagy Szabolcs egyetemi docens Társkonzulens: Dr. Orbán László Készítette: Cserveni-Szücs Réka Keszthely 2013 2 A PONTY (Cyprinus carpio L.) PIKKELYMINTÁZAT ÖRÖKLŐDÉSI MODELLJÉNEK REVÍZIÓJA Értekezés doktori (PhD) fokozat elnyerése érdekében Írta: Cserveni-Szücs Réka Készült a Pannon Egyetem Georgikon Kar Állat-és Agrárkörnyezet-tudományi Doktori Iskolája Keretében Témavezetők: Dr. habil. Bercsényi Miklós egyetemi tanár Dr. Nagy Szabolcs, egyetemi docens Elfogadásra javaslom (igen / nem) …………………………... (aláírás) Elfogadásra javaslom (igen / nem) …………………………... (aláírás) A jelölt a doktori szigorlaton ........%-ot ért el, Az értekezést bírálóként elfogadásra javaslom: Bíráló neve: …........................ …................. igen /nem ………………………. (aláírás) Bíráló neve: …........................ ….................) igen /nem ………………………. (aláírás) ***Bíráló neve: …........................ ….................) igen /nem ………………………. (aláírás) A jelölt az értekezés nyilvános vitáján …..........%-ot ért el. Veszprém/Keszthely, …………………………. a Bíráló Bizottság elnöke A doktori (PhD) oklevél minősítése…................................. ………………………… Az EDHT elnöke 3 Tartalomjegyzék Rövidítések jegyzéke ........................................................................................................ 6 Kivonat .............................................................................................................................. 7 Abstract ............................................................................................................................. 9 Auszug ............................................................................................................................ 10 1. Bevezetés és célkitűzések ........................................................................................... 12 2. Irodalmi áttekintés ...................................................................................................... 15 2.1. A ponty rendszertana ............................................................................................ 16 2.2. A ponty morfológiája /felépítése .......................................................................... 18 2.3. A ponty eredete /elterjedése ................................................................................. 20 2.4. Nemesített és vad ponty változatok ...................................................................... 25 2.4.1. Tájfajták és hibridek kialakulása Magyarországon ....................................... 26 2.4.2. Tájfajták és tenyésztett változatok pikkelyezettsége ..................................... 27 2.5. A ponty evolúciós ugrása – a tetraploid állapot és az ebből adódó lehetőségek .. 28 2.5.1. Egy család- 3 különböző genom méret ......................................................... 28 2.5.2. Genom duplikáció a csontos halakban .......................................................... 29 2.6. Kültakaró és fenotípus .......................................................................................... 31 2.6.1. A pikkely és eredete: ..................................................................................... 33 2.7. A garatfog ............................................................................................................. 35 2.8. A kopoltyú és részei ............................................................................................. 36 2.9. A ponty pikkelymintázatának genetikája anno: ................................................... 37 2.10. Az „s” gén azonosítása ....................................................................................... 41 2.11. A pikkelyezettség kutatásakor eddig vizsgált és az N gén megtalálása szempontjából is jelentősebb szignál transzdukciós kaszkádok ismertetése .............. 43 2.11.1. Fibroblaszt növekedési faktor transzdukciós kaszkád ................................. 44 2.11.2. Az ’ectodysplasin’ anyagcsereútvonal ........................................................ 49 2.11.3. Egyéb, szóba kerülhető kaszkádok .............................................................. 51 3. Anyag és módszer ....................................................................................................... 52 3.1. Az anyapontyok kiválasztása, keresztezésük ....................................................... 52 3.1.1. Magyarországi keresztezések ........................................................................ 52 3.1.2. Szingapúri keresztezések: .............................................................................. 56 3.2. Az ikra felragasztása, termékenyülés számolása: ................................................ 58 4 3.3. A kísérleti halak nevelése: ................................................................................... 59 3.4. A fenotípus kiértékeléséhez szükséges egységes rendszer kialakítása ................ 60 3.5. Kopoltyúfésűk tüskéinek számolása, az úszók szemrevételezése ........................ 62 3.6. Garatfog kiszedése, tisztítása ............................................................................... 63 3.7. Eltérő pikkelyezettségű ponty változatok keresztezéséből származó utódnemzedék növekedésének vizsgálata zárt tartásban ............................................. 64 3.8. Úszóminta gyűjtése és a nukleinsavak izolálása genotipizáláshoz, szekvenáláshoz ..................................................................................................................................... 65 3.9. Szekvenálás és a jelölt gének cDNS-ének összehasonlító vizsgálata .................. 66 3.10. Az alkalmazott statisztikai eljárások és programok ........................................... 66 4. Eredmények és értékelés ............................................................................................. 67 4.1. A keresztezések eredményei Magyarországon: ................................................... 67 4.2. A pikkelyezettséggel összefüggő más paraméterek, szervek vizsgálati eredményei .................................................................................................................. 70 4.2.1. Az úszók, a kopoltyúk, valamint a garatfogak szerkezete ............................. 70 4.2.2. Eltérő pikkelyzettségű ponty változatok keresztezéséből származó utódnemzedék növekedésének kiértékelése ............................................................ 72 4.3. A szingapúri keresztezések kiértékelése .............................................................. 73 4.4. A Magyarországon és Szingapúrban elvégzett 19 keresztezés ............................ 75 4.5. Eltérés a Magyarországon és Szingapúrban végzett keresztezések bőrpontyainál: ..................................................................................................................................... 76 5. Kiegészítő vizsgálatok ................................................................................................ 78 5.1. Genetikai vizsgálatok Szingapúrban .................................................................... 78 A ponty fgfr újabb paralógjainak felfedezése .......................................................... 80 5.2. Magyarországi kiegészítő vizsgálatok ................................................................. 80 5.2.1. A különböző fenotípusú pontyok pikkelyeinek alaktani vizsgálata .............. 80 5.2.2. „Karcolásos kísérlet” ..................................................................................... 81 6. Következtetések és Javaslatok .................................................................................... 83 7. Összefoglalás .............................................................................................................. 88 Irodalomjegyzék ............................................................................................................. 90 Tézispontok ................................................................................................................... 108 Thesis Points ................................................................................................................. 109 Köszönetnyilvánítás ...................................................................................................... 110 Tudományos közlemények jegyzéke ............................................................................ 111 Melléklet ....................................................................................................................... 114 5 Rövidítések jegyzéke DAG: diacilglicerol DDC: (Duplication- Degeneration- Complementation) duplikáció-degeneráció-komplementáció modell EDA: ectodysplasin-A EDAR: ectodysplasin-A receptor EDARADD: EDAR-(associated death domain) kapcsolódó haláldomén FCR: takarmányhasznosítási érték FGF: Fibroblast growth factor: Fibroblast növekedési faktor Fgfr1: Fibroblast growth factor receptor 1: Fibroblast növekedési faktor receptor 1 HAKI: Haltenyésztési Kutató Intézet (Szarvas) Ic: (incomplete scaled) nem teljesen pikkelyes IKK: IkappaB kináz (IKK) IP3: inozitol 1,4,5-trisfoszfát Ir: (irregular) szabálytalan Li: (linear) oldalvonalsoros, keretes m:
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