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SZENT ISTVÁN EGYETEM Kertészettudományi Kar SZENT ISTVÁN EGYETEM Kertészettudományi Kar SORBUS FAJKELETKEZÉS TRIPARENTÁLIS HIBRIDIZÁCIÓVAL A KELET- ÉS DÉLKELET- EURÓPAI TÉRSÉGBEN (Nothosubgenus Triparens) Doktori (PhD) értekezés Németh Csaba BUDAPEST 2019 A doktori iskola megnevezése: Kertészettudományi Doktori Iskola tudományága: Növénytermesztési és kertészeti tudományok vezetője: Zámboriné Dr. Németh Éva egyetemi tanár, DSc Szent István Egyetem, Kertészettudományi Kar, Gyógy- és Aromanövények Tanszék Témavezető: Dr. Höhn Mária egyetemi docens, CSc Szent István Egyetem, Kertészettudományi Kar, Növénytani Tanszék és Soroksári Botanikus Kert A jelölt a Szent István 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. .................................................. .................................................. Az iskolavezető jóváhagyása A témavezető jóváhagyása 2 Édesanyám emlékének. 3 4 TARTALOMJEGYZÉK RÖVIDÍTÉSEK JEGYZÉKE .......................................................................................................... 7 1. BEVEZETÉS ÉS CÉLKITŰZÉS .................................................................................................. 9 2. IRODALMI ÁTTEKINTÉS ..................................................................................................... 11 2.1. A Sorbus nemzetség taxonómiai vonatkozásai ..................................................................... 11 2.2. Az európai Sorbus alnemzetségek növényföldrajzi vonatkozásai ......................................... 13 2.3. A fajkeletkezés mozgatórugói a Sorbus nemzetségen belül .................................................. 15 2.3.1. Hibridizáció .............................................................................................................................................. 16 2.3.1.1. Pollendonáció az endospermium képzés során ............................................................................... 16 2.3.1.2. Pollendonáció a hibridizációs fajkeletkezés során ........................................................................... 17 2.3.2. Poliploidizáció .......................................................................................................................................... 19 2.3.3. Reprodukciós mód ................................................................................................................................... 19 2.4. Hármas hibridizáció a növényvilágban ................................................................................ 20 2.5. Hármas hibridizáció a Sorbus nemzetségben ....................................................................... 20 2.5.1. Nothosubg. Chamsoraria (Subg. Chamaemespilus × Subg. Sorbus × Subg. Aria) .................................... 21 2.5.2. Nothosubg Triparens (Subg. Sorbus × Subg. Aria × Subg. Torminaria). ................................................... 21 2.5.2.1. Bizonyítottan Triparens fajok ........................................................................................................... 21 2.5.2.2. Feltételezett Triparens fajok ............................................................................................................ 23 2.6. Az európai Sorbus alnemzetségek taxonómiai markerei ...................................................... 26 2.6.1. Morfológiai markerek .............................................................................................................................. 26 2.6.2. Többváltozós morfometriai vizsgálatok................................................................................................... 31 2.6.2. Biokémiai markerek ................................................................................................................................. 32 2.6.3. Molekuláris markerek .............................................................................................................................. 33 2.6.3.1. Izoenzim markerek .......................................................................................................................... 33 2.6.3.2. Mitokondriális DNS markerek .......................................................................................................... 34 2.6.3.3. Kloroplasztisz DNS markerek ........................................................................................................... 34 2.6.3.4. Nukleáris DNS markerek .................................................................................................................. 35 2.6.3.5. Egyéb molekuláris markerek ............................................................................................................ 36 2.6.4. Citometriai markerek ............................................................................................................................... 36 3. ANYAG ÉS MÓDSZER ........................................................................................................ 39 3.1. Terepi mintavételezés .......................................................................................................... 39 3.2. Morfológiai vizsgálatok ....................................................................................................... 39 3.2.1. Herbáriumi kutatások .............................................................................................................................. 39 3.2.2. Morfometriai vizsgálatok (PCA) ............................................................................................................... 42 3.3. Biokémiai vizsgálatok (HPLC-ESI-qTOFMS) ..................................................................... 43 3.3.1. Minta előkészítés ..................................................................................................................................... 43 3.3.2. Kromatográfiás elválasztás ...................................................................................................................... 43 3.3.3. HPLC-ESI-qTOFMS analízis ....................................................................................................................... 43 3.4. Molekuláris vizsgálatok (cpSSR) ......................................................................................... 44 3.4.1. DNS-kivonás ............................................................................................................................................. 44 3.4.2. Fragmens amplifikáció PCR-rel ................................................................................................................ 45 3.5. Áramlási citometriás vizsgálatok (FCM, FCSS) ................................................................... 46 3.5.1. Ploidia szint meghatározás (FCM) ........................................................................................................... 46 3.5.2. Reprodukciós mód meghatározás (FCSS) ................................................................................................ 46 4. EREDMÉNYEK ÉS MEGVITATÁSUK..................................................................................... 48 4.1. Morfológiai vizsgálatok ....................................................................................................... 48 4.1.1. Herbáriumi kutatások .............................................................................................................................. 48 5 4.2. Biokémiai vizsgálatok (HPLC-ESI-qTOFMS) ..................................................................... 49 4.2.1. Flavon O-glükozid (Apigenin O-glükuronsav) .......................................................................................... 49 4.2.2. Flavon C-glükozid (Apigenin-8-C-glükozid, vitexin) .................................................................................. 50 4.3. Molekuláris vizsgálatok (cpSSR) ......................................................................................... 53 4.4. Áramlási citometriás vizsgálatok (FCM, FCSS) ................................................................... 55 4.4.1. Ploidia szint meghatározás (FCM) ........................................................................................................... 55 4.4.2. Reprodukciós mód meghatározás (FCSS) ................................................................................................ 56 4.5. A Sorbus dacica s.l. taxonómiai revíziója ............................................................................. 58 4.5.1. A Sorbus dacica s.l. a szakirodalmi forrásokban ...................................................................................... 58 4.5.2. A Sorbus dacica s.l. a herbáriumi forrásokban ........................................................................................ 60 4.5.3. A Sorbus dacica s.l. a terepen, földrajzi elterjedés .................................................................................. 62 4.5.4. A Sorbus dacica s.l. morfológiai variabilitása ........................................................................................... 63 4.5.5. A Sorbus dacica s.l. kemotaxonómiai variabilitása .................................................................................. 65 4.5.6. A Sorbus dacica s.l. kloroplasztisz variabilitása ....................................................................................... 66 4.5.7. A Sorbus dacica s.l. citotípus variabilitása ..............................................................................................
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