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Mendelova Univerzita V Brně Hybridizace V Rámci Rodu Carex A

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Mendelova Univerzita V Brně Hybridizace V Rámci Rodu Carex A Mendelova univerzita v Brn ě Lesnická a d řeva řská fakulta Ústav lesnické botaniky, dendrologie a geobiocenologie Hybridizace v rámci rodu Carex a zp ůsoby jejího testování Diserta ční práce 2016 Petra Veselá Čestné prohlášení Prohlašuji, že jsem práci: „Hybridizace v rámci rodu Carex a zp ůsoby jejího testování“ vypracovala samostatn ě a veškeré použité prameny a informace uvádím v seznamu použité literatury. Nesouhlasím, aby moje práce byla zve řejn ěna v plném rozsahu v souladu s § 47b zákona č. 111/1998 Sb.,o vysokých školách ve zn ění pozd ějších p ředpis ů a v souladu s platnou Sm ěrnicí o zve řej ňování vysokoškolských záv ěre čných prací. Jsem si v ědoma, že se na moji práci vztahuje zákon č. 121/2000 Sb., autorský zákon, a že Mendelova univerzita v Brn ě má právo na uzav ření licen ční smlouvy a užití této práce jako školního díla podle § 60 odst. 1 autorského zákona. Dále se zavazuji, že p řed sepsáním licen ční smlouvy o využití díla jinou osobou (subjektem) si vyžádám písemné stanovisko univerzity, že p ředm ětná licen ční smlouva není v rozporu s oprávn ěnými zájmy univerzity, a zavazuji se uhradit případný p řísp ěvek na úhradu náklad ů spojených se vznikem díla, a to až do jejich skute čné výše. V Brn ě dne:……………………….. …………………………………………………….. podpis POD ĚKOVÁNÍ Zpracovaná diserta ční práce byla finan čně podpo řena z prost ředk ů specifického vysokoškolského výzkumu prost řednictvím projektu IGA LDF č. 11/2013 a LDF_VP_2015024. Své pod ěkování chci také v ěnovat Dr. Ing. Jaroslavu Mrá čkovi za podporu v doktorském studiu, koleg ům, které se podílely na mém profesním rozvoji a rodin ě za morální podporu. Petra Veselá Název: Hybridizace v rámci rodu Carex a zp ůsoby jejího testování Abstrakt K objasn ění široké škály p řírodních vztah ů, jak vnitrodruhových, tak i mezidruhových, může skv ěle posloužit rod Carex. Jedná se o nejv ětší rod čeledi Cyperacea a jeden z celosv ětov ě nejrozší řen ějších rod ů, který je také zajímavý n ěkolika specifickými rysy. Krom ě již zmín ěné druhové diverzity je pro tento rod charakteristická zna čná vnitrodruhová variabilita a také relativn ě častá hybridizace. Dále jsou to cytogenetická specifika jako je p řítomnost holocentrických chromozom ů, což je spojeno s agmatoploidií a symplodií či invertovaná meióza. Cílem této práce bylo objasnit p říčiny morfologické variability n ěkolika domn ělých hybridních jedinc ů vzniklých k řížením r ůzných kombinací parentálních druh ů pomocí molekulárních marker ů. Jako vhodné markery byly zvoleny AFLP a mikrosatelitové markery, nebo ť mají dostate čnou rozlišovací schopnost mezi blízce p říbuznými druhy a vysokou informa ční hodnotu. Mezi analyzované kombinace rodi čovských druh ů byly C. flacca subsp. flacca a C. tomentosa , C. acutiformis a C. nigra , C. caryophyllea a C. fritschi , C. paniculata a C. echinata , C. paniculata a C. appropinquata . Ve v ětšin ě p řípad ů bylo dosaženo zna čné genetické diferenciace mezi páry taxon ů zjišt ěné pomocí Neiových genetických vzdáleností, ale výjimku tvo řila kombinace C. paniculata a C. appropinquata , což mohlo být zap říčin ěno velmi blízkým vztahem t ěchto taxon ů. Nicmén ě z těchto p ěti kombinací byla pouze u C. paniculata × echinata detekována genetická intermediárnost na základ ě molekulárních marker ů. Získané výsledky na jednu stranu poukazují na p řítomnost mezidruhové hybridizace u n ěkterých kombinací. Ale na stranu druhou, domn ělá morfologická intermediárnost mezi dv ěma druhy m ůže být p řipsána dosud nedetekované vnitrodruhové morfologické variabilit ě. Tato práce dokládá zna čný význam, a to nejenom ve sporných p řípadech, molekulárních marker ů, které dokáží oz řejmit genetické pozadí těchto taxon ů. Klí čová slova : AFLP, Carex , hybridizace, mikrosatelitové markery Title: Hybridisation within the genus Carex and the methods for its testing Abstract To clarify the wide range of natural relationships, intra- as well as interspecific, may serve the genus Carex . It is the largest genus of the family Cyperacea and one of the most widespread genera worldwide, which is also interesting several specific features. Instead the above mentioned species diversity, it is for the genus characteristic considerable intraspecific variability and relatively frequent hybridization. Furthermore, there are also cytogenetic features, such as the presence of holocentric chromosomes, which is associated with agmatoploidy and symplody or inverted meiosis. The aim of this study was to clarify the causes of morphological variability of several putative hybrids resulting from cross of different combinations of the parental taxa using molecular markers. As suitable markers were chosen AFLP and microsatellite markers, because they have sufficient resolution power between closely related species and also high information value. Among the analysed combinations of parental species were C. flacca subsp. flacca and C. tomentosa , C. acutiformis and C. nigra , C. caryophyllea and C. fritschii , C. paniculata and C. echinata , C. paniculata and C. appropinquata . In most cases, it was achieved significant genetic differentiation between taxon pairs according to Nei’s genetic distances, with the exception of combination C. paniculata and C. appropinquata , which could be due to the very close relationship of these taxa. However, from these five combinations it was only in C. paniculata × echinata detected genetic intermediary based on molecular markers. The obtained results indicated the presence of interspecific hybridization. On the other hand, putative morphological intermediary between two species can be ascribed to the yet undetected intraspecific morphological variability. This work also gives the evidence of considerable importance of molecular markers that can elucidate the genetic background of these taxa. Keywords : AFLP, Carex , hybridisation, microsatellite markers OBSAH 1. ÚVOD ................................................................................................................................ 1 2. CÍLE PRÁCE .................................................................................................................... 2 3. STRUKTURA PRÁCE ..................................................................................................... 3 4. ÚVOD DO PROBLEMATIKY ........................................................................................ 4 4.2 Hybridizace u rostlin .................................................................................................... 4 4.3 Rod Carex a jeho obecné vymezení ............................................................................ 8 4.4 Markery ...................................................................................................................... 22 4.4.1 Polymorfní náhodn ě amplifikovaná DNA (RAPD) ............................................ 24 4.4.2 Polymorfismus délky amplifikovaných fragment ů (AFLP)................................ 25 4.4.3 Mikrosatelitové markery ..................................................................................... 26 5. LITERATURA ................................................................................................................ 28 Přílohy I. Optimisation of AFLP for extremely large genomes over 70 Gb II. Are there hybrids between Carex flacca and C. tomentosa in the Czech Republic and Slovakia? III. Morphological and genetic variation of several putative Carex hybrids 1. ÚVOD Molekulární biologie p ředstavuje dynamicky se rozvíjející v ědní obor a nachází široké spektrum uplatn ění p ři řešení nejr ůzn ějších problematik. Význam molekulární biologie se stále zvyšuje a její využití ve stále v ětší mí ře přesahuje do řady jiných disciplín, jako jsou nap říklad medicína či zem ědělství a lesnictví. Pot řeba analýzy stále v ětších objem ů dat sou časn ě při stále se rozši řujícím poli působnosti vedou nejen k vývoji nových metod zpracování a analýzy dat, jaké představují nap říklad metody NGS (Next Generation Sequencing). Na stranu druhou své uplatn ění nacházejí i metody konven ční, jakými je sekvenování jednotlivých lokus ů, nebo nap říklad metoda AFLP, která představuje metodu tak řka univerzální a navíc využitelnou i pro druhy, u nichž nejsou prozatím k dispozici specifické DNA markery (Vos et al. 1995; Qi a Lindhout, 1997). Tato práce je zaměř ená na využití molekulárních marker ů p ři determinace mezidruhové hybridizace v rodu Carex, nebo ť práv ě pro tento rod je hybridizace relativn ě často popisovaným jevem (Ford et al. 1993; Smith a Waterway, 2008; Volkova et al. 2008; Korpelainen et al. 2010), ale nemalá část publikací se opírá pouze o markery morfologické (Catling, 1993; Blackstock a Ashton, 2010; Wieclaw a Koopman, 2013; Bergeron a Pellerin, 2014; Wieclaw a Wilhelm, 2014) a jak je patrné z výsledk ů této i dalších prací (Escudero et al. 2014, Řepka et al. 2014), nevede tento postup vždy ke správným záv ěrům. Na základ ě t ěchto indicií je možné předpokládat, že hybridizace v rámci rodu Carex je nadhodnocená a naopak podhodnocená je vnitrodruhová variabilita taxon ů. Část práce je také v ěnována optimalizaci AFLP, která – ač s četným uplatn ěním – má také své nedostatky, jakým je nap říklad velikostní homoplasie. 1 2. CÍLE PRÁCE Rod Carex je charakteristický zna čnou vnitrodruhovou variabilitou a relativn ě častou hybridizace a tyto vlastnosti dále komplikují determina a taxonomicko klasifikaci. Cílem této práce bylo objasnit genetické pozadí morfologické variability n ěkolika domn ělých hybridních jedinc ů rodu Carex pomocí vhodných molekulárních
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