Kovaalga Fajok Filogenetikai És Morfológiai Diverzitása

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Kovaalga Fajok Filogenetikai És Morfológiai Diverzitása Duleba Mónika Kovaalga fajok filogenetikai és morfológiai diverzitása Doktori értekezés Témavezető: Dr. Ács Éva, DSc tudományos tanácsadó MTA Ökológiai Kutatóközpont, Duna-kutató Intézet Környezettudományi Doktori Környezetbiológia Doktori Iskola Program Iskolavezető: Dr. Jánosi Imre, DSc Programvezető: Dr. Ács Éva, DSc Készült: Eötvös Loránd Tudományegyetem, Budapest 2017 1 Tartalom Rövidítések jegyzéke .................................................................................................................. 4 1. Bevezetés ................................................................................................................................ 6 2. Célkitűzések ........................................................................................................................... 8 3. Irodalmi áttekintés .................................................................................................................. 9 3.1. A molekuláris taxonómiai módszerek szerepe a kovaalgák taxonómiájában .............................. 9 3.2. A Thalassiosirales rend evolúciója ............................................................................................. 15 3.3. Fajfogalmak a kovaalga taxonómiában ...................................................................................... 22 3.4. A fajok DNS vonalkódja ............................................................................................................ 25 3.5. Molekuláris markerek a kovaalga taxonómiában és filogenetikában ......................................... 28 3.5.1. A riboszómális RNS ............................................................................................................ 28 3.5.2. A sig1 .................................................................................................................................. 34 3.5.3. Mitokondriális markerek ..................................................................................................... 34 3.5.4. Kloroplasztisz markerek ...................................................................................................... 34 3.6. A morfológiai és a genetikai variabilitás összekötése ................................................................ 38 4. Anyag és módszer ................................................................................................................ 41 4.1. Mintavétel ................................................................................................................................... 41 4.2. A környezeti változók mérése .................................................................................................... 42 4.3. Biomassza meghatározás és morfológiai vizsgálat .................................................................... 43 4.4. Morfológiai vizsgálat ................................................................................................................. 43 4.5. A Cyclotella ocellata/Cyclotella comensis komplex morfometriai vizsgálata ........................... 44 4.6. Molekuláris vizsgálatok ............................................................................................................. 45 4.6.1. DNS-kivonás ....................................................................................................................... 45 4.6.2. Primertervezés és polimeráz láncreakció ............................................................................ 47 4.6.3. A szekvenciák elemzése: filogenetika és genetikai távolság ............................................... 50 4.7. Statisztikai elemzések ................................................................................................................. 53 5. A Skeletonema potamos vizsgálata ...................................................................................... 55 5.1. Bevezetés .................................................................................................................................... 55 5.1.1. A Skeletonema nemzetség és a Skeletonema potamos ........................................................ 55 5.1.2. A Skeletonema potamos elterjedése és előfordulása a hazai vizekben ................................ 56 5.2. Eredmények ................................................................................................................................ 57 5.2.1. Morfológiai jellemzés .......................................................................................................... 57 5.2.2. A primerek specificitása ...................................................................................................... 58 5.2.3. Filogenetikai elemzés .......................................................................................................... 59 5.2.4. A Skeletonema potamos földrajzi elterjedése ...................................................................... 63 2 5.2.5. A Skeletonema potamos mennyiségének hosszú távú változása a Dunában a környezeti változók alakulásával összefüggésben .......................................................................................... 63 5.3. Diszkusszió ................................................................................................................................. 66 5.3.1. A Skeletonema potamos morfológiája ................................................................................. 66 5.3.2. A Skeletonema potamos filogenetikai helyzete ................................................................... 66 5.3.3. A Skeletonema potamos földrajzi elterjedése ...................................................................... 67 5.3.4. A Skeletonema potamos és a Duna környezeti változásai közti kapcsolat .......................... 67 6. A Cyclotella ocellataval és rokonsági körével kapcsolatos vizsgálatok .............................. 71 6.1. Bevezetés .................................................................................................................................... 71 6.1.1. A Cyclotella ocellata előfordulása és morfológiai variabilitása, a Cyclotella ocellata/Cyclotella comensis komplex .......................................................................................... 71 6.2. Eredmények ................................................................................................................................ 73 6.2.1. A morfológiai és morfometriai vizsgálat ............................................................................. 73 6.2.2. A morfológiai jellemvonások kapcsolata a környezeti változókkal .................................... 82 6.2.3. A Cyclotella ocellata komplexhez tartozó minták DNS-einek szekvencia meghatározása 83 6.2.4. Genetikai divergencia .......................................................................................................... 85 6.2.5. Filogenetika ......................................................................................................................... 88 6.2.6. A morfometriai és a filogenetikai adatok összevetése ......................................................... 89 6.3. Diszkusszió ................................................................................................................................. 90 6.3.1. A Cyclotella ocellata komplex morfológiai és morfometriai vizsgálata ............................. 90 6.3.2. A filogenetikai divergencia és a DNS vonalkódok használata ............................................ 93 6.3.3. A filogenetikai összefüggések a Cyclotella ocellata/Cyclotella comensis komplexben ..... 96 7. A Cyclotella ocellata filogenetikai vizsgálata – a Pantocsekiella nemzetség ..................... 98 7.1. Bevezetés .................................................................................................................................... 98 7.1.1. A Cyclotella nemzetség ....................................................................................................... 98 7.2. Eredmények .............................................................................................................................. 100 7.2.1. Morfológiai vizsgálat......................................................................................................... 100 7.2.2. A vizsgált fajok filogenetikai vizsgálata ........................................................................... 101 7.3. Diszkusszió ............................................................................................................................... 106 8. Összefoglalás ...................................................................................................................... 116 9. Summary ............................................................................................................................ 119 10. Köszönetnyilvánítás ......................................................................................................... 122 11. Irodalom ........................................................................................................................... 123 12. Függelék ................................................................................................................................ I 3 Rövidítések jegyzéke Ahol a rövidítés angol kifejezésből származik, ott ezt a kifejezést zárójelben adtuk meg. 18S rDNS – riboszóma kis alegység (18S) KOI – kémiai oxigénigény rRNS gén l – pervalváris tengely hossza 28S rDNS – riboszóma nagy alegység (28S) LSU – nagy alegység (large subunit) rRNS gén MEGA – Molecular Evolution
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