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A Jósika-Orgona (Syringa Josikaea Jacq

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A Jósika-Orgona (Syringa Josikaea Jacq Doktori (Ph.D.) értekezés A JÓSIKA-ORGONA (SYRINGA JOSIKAEA JACQ. FIL. EX RCHB.) AKTUÁLIS ÉS TÖRTÉNETI BIOGEOGRÁFIÁJA Lendvay Bertalan Kertészettudományi Doktori Iskola Budapesti Corvinus Egyetem Genetika és Növénynemesítés Tanszék Budapest 2014 A doktori iskola megnevezése: Kertészettudományi Doktori Iskola tudományága: Növénytermesztési és kertészeti tudományok vezetője: Dr. Tóth Magdolna egyetemi tanár, DSc Budapesti Corvinus Egyetem, Kertészettudományi Kar, Gyümölcstermő Növények Tanszék Témavezetők: Dr. Höhn Mária egyetemi docens, CSc Budapesti Corvinus Egyetem, Kertészettudományi Kar, Növénytani Tanszék és Soroksári Botanikus Kert Dr. Pedryc Andrzej egyetemi tanár, DSc Budapesti Corvinus Egyetem, Kertészettudományi Kar, Genetika és Növénynemesítés Tanszék A jelölt a Budapesti Corvinus Egyetem Doktori Szabályzatában előírt valamennyi feltételnek eleget tett, az értekezés műhelyvitájában elhangzott észrevételeket és javaslatokat az értekezés átdolgozásakor figyelembe vette, azért az értekezés védési eljárásra bocsátható. ……………………………… ……………………………… Dr. Tóth Magdolna Dr. Höhn Mária ……………………………… Dr. Pedryc Andrzej A doktori iskola vezetőjének A témavezetők jóváhagyása jóváhagyása 2 A Budapesti Corvinus Egyetem Élettudományi Területi Doktori Tanácsának 2014.03.18-i határozatában a nyilvános vita lefolytatására az alábbi bíráló Bizottságot jelölte ki: BÍRÁLÓ BIZOTTSÁG: Elnöke Palkovics László, DSc Tagjai Bisztray György, PhD Deák Tamás, PhD Csiky János, PhD Pócs Tamás, MHAS Opponensek Halász Júlia, PhD Kovács J. Attila, CSc Titkár Deák Tamás, PhD 3 Tartalomjegyzék Rövidítések jegyzéke ........................................................................................ 6 Táblázatok jegyzéke......................................................................................... 7 Ábrák jegyzéke................................................................................................. 7 1. Bevezetés ....................................................................................................... 8 2. Célkitűzés.................................................................................................... 10 3. Irodalmi áttekintés ..................................................................................... 11 3.1. A Syringa josikaea bemutatása ........................................................................................... 11 3.1.1. A Syringa josikaea alaktani leírása .............................................................................. 11 3.1.2. A Syringa josikaea élőhelye......................................................................................... 11 3.1.2. A Syringa josikaea rendszertani helyzete...................................................................... 12 3.1.3. A Syringa josikaea szűkebben vett rokonsága.............................................................. 14 3.2. Fosszilis Syringa maradványok Európából és elméletek a Syringa josikaea származásával kapcsolatban ............................................................................................................................. 15 3.3. Eurázsia flórájának alakulása a Kainozoikumban, eurázsiai diszjunkciók és Tercier maradványfajok Európában....................................................................................................... 17 3.4. Populációgenetikai vizsgálati módszerek és vizsgálatok a Syringa josikaea rokonainál....... 20 3.4.1. Nukleotid sorrenden alapuló DNS markerek ................................................................ 21 3.4.2. Hosszpolimorfizmuson alapuló DNS markerek............................................................ 22 3.5. A Syringa josikaea Erdélyi-szigethegységi elterjedésének szakirodalma............................. 23 3.5.1. A Syringa josikaea felfedezése az Erdélyi-szigethegységben ....................................... 24 3.5.2. Az első előfodulási adatok 1830-tól 1880-ig ................................................................ 25 3.5.3. Az elterjedési terület feltárása az 1880-as években....................................................... 26 3.5.4. 1890-1908: áttekintő tanulmányok az addig megismert elterjedésről............................ 28 3.5.5. Az elterjedés további pontosítása Blattny Tibor nyomán 1909-től 1913-ig ................... 29 3.5.6. Fekete Lajos és Blattny Tibor – a korábbról ismert elterjedési adatok és az újabb kutatási eredmények szintézise, 1913 ................................................................................................. 31 3.5.7. Az I. világháborútól napjainkig keletkezett szakirodalom............................................. 33 3.5.8. A munkám kezdetéig elfogadott Syringa josikaea elterjedési adatok az Erdélyi- szigethegységben................................................................................................................... 34 3.6. A Syringa josikaea elterjedési adatai az Ukrajnai-Kárpátokban........................................... 37 4. Anyag és módszer....................................................................................... 40 4.1. A Syringa josikaea Erdélyi-szigethegységi elterjedésének felmérése .................................. 40 4.2. Mintagyűjtés és DNS kivonás............................................................................................. 40 4.3. Molekuláris módszerek....................................................................................................... 41 4.3.1. Fajon belül variábilis DNS szakaszok keresése ............................................................ 41 4.3.2.Nukleáris mikroszatellit markerek tesztelése és fejlesztése............................................ 46 4.3.2.1. Olea és Ligustrum mikroszatellit markerek használhatósága a Syringa josikaea esetében ................................................................................................................................ 46 4.3.2.2. De novo mikroszatellit marker fejlesztés................................................................... 47 4.4. Az adatelemzés módszerei.................................................................................................. 50 4.4.1. A Syringa josikaea divergencia idejének kiszámítása molekuláris óra analízissel......... 50 4.4.2. Szekvencia variabilitás a Syringa josikaea populációk közt ......................................... 52 4.4.3. Mikroszatellit markerek tesztelése és fejlesztése .......................................................... 53 4.4.4. A mikroszatellit adatok kiértékelése és adatelemzése ................................................... 53 4.4.4.1. Az egy klónhoz tartozó minták összevonása.............................................................. 53 4 4.4.4.2. A populációk genetikai diverzitása............................................................................ 54 4.4.4.3. A populációk genetikai különbségei földrajzi helyzetük figyelembevételével............ 55 4.4.4.4. A genetikai és földrajzi távolságok korrelációja ........................................................ 56 5. Eredmények................................................................................................ 57 5.1. A Syringa josikaea szakirodalomban jelzett állományainak felkutatása az Erdélyi- szigethegységben ...................................................................................................................... 57 5.2. Divergencia idő meghatározása .......................................................................................... 64 5.3. DNS szekvenciák változékonyságának tesztelése a Syringa josikaea populációi közt.......... 65 5.4. A variábilis DNS szakaszok fajon belüli variabilitása és eloszlása ...................................... 66 5.5. Mikroszatellit markerek tesztelése és tervezése................................................................... 69 5.5.1. Olea és Ligustrum mikroszatellit markerek használhatósága a Syringa josikaea esetében .............................................................................................................................................. 69 5.5.2. De novo mikroszatellit markertervezés ........................................................................ 70 5.5.3. A mikroszatellit markerek jellemzése........................................................................... 70 5.6. A populációk mikroszatellit elemzése................................................................................. 72 6. Következtetések.......................................................................................... 78 6.1. A Syringa josikaea aktuális elterjedése az Erdélyi-szigethegységben .................................. 78 6.2. A Syringa josikaea populációk veszélyeztető tényezői az Erdélyi-szigethegységben........... 78 6.3. A Syringa josikaea divergencia ideje .................................................................................. 79 6.4. Kismértékű szekvencia variabilitás a Syringa josikaea populációi közt............................... 81 6.5. A riboszomális lókuszok típusai és leszármazása................................................................ 81 6.5. Mikroszatellit diverzitás ..................................................................................................... 82 6.5.1. Mikroszatellit markerek tesztelése és fejlesztése .......................................................... 82 6.5.2. A Syringa josikaea mikroszatellit diverzitása............................................................... 83 6.6. A Syringa josikaea két diszjunkt areajának
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