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Bakalářská Práce Masarykova univerzita Přírodov ědecká fakulta Ústav botaniky a zoologie Korelace velikosti pr ůduch ů a genomických parametr ů kapra ďorost ů Bakalá řská práce Michaela Burešová Brno 2011 Vedoucí bakalá řské práce: Mgr. Petr Šmarda, Ph.D. 2 Prohlášení „Souhlasím s uložením této bakalá řské práce v knihovn ě Ústavu botaniky a zoologie P řF MU v Brn ě, p řípadn ě v jiné knihovn ě MU, s jejím ve řejným p ůjčováním a využitím pro vědecké, vzd ělávací nebo jiné ve řejn ě prosp ěšné ú čely, a to za p ředpokladu, že p řevzaté informace budou řádn ě citovány a nebudou využívány komer čně.“ 3 4 Pod ěkování Na prvním míst ě pat ří jist ě mé velké pod ěkování panu Dr. Petru Šmardovi 1) , vedoucímu této bakalá řské práce, za to, že mi velmi ochotn ě poskytoval časté odborné konzultace a trp ěliv ě mi vše vysv ětloval; nemalý dík pat ří také mému otci, panu doc. Petru Burešovi 1) , který mi byl p ři této práci inspirací a velkou duševní oporou; ob ěma jsem zavázána za přečtení celé práce ve finální fázi a za množství kritických p řipomínek, které mi usnadnily zejména interpretaci a diskuzi získaných výsledk ů. Za cytometrická m ěř ení, p ředcházející této studii, vd ěč ím Mgr. Lucii Horové 1) a Mgr. Klá ře Helánové 1) , bez t ěchto základních dat by nebylo možné studii v tomto rozsahu uskute čnit. Rovn ěž d ěkuji sb ěratel ům analyzovaných vzork ů, kterými jsou krom ě výše jmenovaných Dr. Lubomír Adamec 2) , doc. Vít Grulich 1) , Mgr. Ivana Hralová 1) a Dr. Olga Rotreklová 1) . Za determinaci a revizi ur čení dokladového materiálu d ěkuji panu Hermannu Esserovi 3) a doc. Vladimíru Řeho řkovi 1) . Finan ční zázemí práce tvo řily projekty LC06073 a MSM0021622416 poskytnuté Ministerstvem školství, mládeže a t ělovýchovy; p řístrojové vybavení a software poskytly laborato ř pr ůtokové cytometrie rostlin a karyologická laborato ř rostlin Ústavu botaniky a zoologie Přírodov ědecké fakulty Masarykovy univerzity v Brn ě; rostlinný materiál z v ětší části poskytly botanické zahrady Masarykovy univerzity v Brn ě, Mendelovy univerzity v Brn ě, Univerzity ve Vídni, Univerzity v Regensburgu, dále botanické zahrady ve Vídni- Schönbrunnu, Mnichov ě-Nymphenburgu, v Bormiu, Praze-Tróji a také Sbírka vodních a mok řadních rostlin Botanického ústavu AV ČR v T řeboni. Za finan ční, p řístrojové i materiálové zázemí všem t ěmto institucím d ěkuji. 1) Ústav botaniky a zoologie, P řírodov ědecké fakulty Masarykovy univerzity v Brn ě 2) Botanický ústav Akademie v ěd České republiky v Pr ůhonicích, pracovišt ě T řebo ň 3) Pteridologické odd ělení skleníku botanické zahrady Mnichov (München, Nymphenburg) 5 6 Abstrakt V této práci se zabývám vztahem mezi obsahem DNA, genomickým obsahem GC bazí a délkou sv ěracích bun ěk pr ůduch ů u recentních kapra ďorost ů ( Monilophyta ). Tento vztah byl zatím studován p ředevším u semenných rostlin, kde byla zjišt ěna pozitivní korelace mezi obsahem DNA a velikostí pr ůduch ů. U kapra ďorost ů toto ješt ě zkoumáno nebylo. Mikroreliéfovou metodou byla analyzována velikost pr ůduch ů u celkem 115 druh ů kapra ďorost ů se známým obsahem DNA a GC bazí. Výb ěr druhů pokrývá většinu hlavních fylogenetických linií recentních kapra ďorost ů. Vzájemné vztahy genomických a anatomických znak ů byly testovány b ěžnými statistickými metodami a pomocí fylogeneticky nezávislých kontrast ů. Kapra ďorosty mají velkou variabilitu jak v obsahu DNA, tak ve velikosti pr ůduch ů. Pr ůměrné hodnoty t ěchto dvou znak ů jsou u kapra ďorost ů vyšší než u krytosemenných rostlin. Podíl GC bazí v genomu kapra ďorost ů je podobný hodnotám zjišt ěným u ostatních skupin vyšších rostlin. Mezi obsahem DNA a velikostí pr ůduch ů byla pomocí fylogeneticky nezávislých kontrast ů prokázána pozitivní korelace; ostatní vztahy mezi zkoumanými parametry žádnou korelaci nevykazovaly. Vztah mezi obsahem DNA a velikostí pr ůduch ů u kapra ďorost ů má tedy stejný trend jako u nahosemenných a krytosemenných rostlin. Je proto možné, že jak u krytosemenných rostlin, tak u kapradin by vývoj velikosti genomu v prehistorických dobách mohl být zkoumán pomocí m ěř ení pr ůduch ů u fosilních doklad ů vym řelých druh ů. Klí čová slova: fylogeneticky nezávislé kontrasty, genomický obsah GC, kapradiny, kapra ďorosty, Monilophyta , obsah DNA, pr ůduchy, velikost genomu, velikost pr ůduch ů 7 Abstract Correlation of stomatal size and genomic parameters in Monilophytes In this thesis I studied relationship between DNA content, genomic GC content and guard cell length in extant ferns ( Monilophyta ). This relationship has been studied particularly in seed plants where positive correlation between DNA content and stomatal length was documented. However, this relationship has never been tested in ferns. Stomatal length of 115 species with previously known genome size and GC content was measured using microrelief reprints of epidermal surface. The sampling representatively covers the complete phylogenetic tree of extant ferns. The relationships of investigated features were tested using common statistic methods and phylogenetically independent contrasts. Ferns show high variation in DNA content and guard cell length; mean values of these traits are higher than in angiosperms. GC content in fern genomes is similar to other groups of vascular plants. Both correlation analysis and analysis of phylogenetically independent contrasts revealed that DNA content in ferns is positively correlated with guard cell length. No other significant correlations were found between other studied traits. Therefore the relationship between DNA content and guard cell length seems universal both in ferns and in angiosperms. This finding indicates that both in ferns and angiosperms, early evolution of plant genome size might be studied based on the measurement of stomatal size in fossil species. Key words: DNA amount, ferns, genome size, genomic GC content, guard cells, Monilophyta , phylogenetically independent contrasts, pteridophytes, stomatal size 8 Obsah 1. Úvod ........................................................................................................................... 11 2. Kapra ďorosty: fylogenetické postavení, životní cyklus a polyploidie ....................... 13 2.1. Fylogenetické postavení a hlavní vývojové linie .............................................. 13 2.2. Životní cyklus .................................................................................................... 14 2.3. Polyploidie u kapra ďorost ů ............................................................................... 15 3. Pr ůduchy ..................................................................................................................... 21 4. Velikost genomu ......................................................................................................... 27 5. Pom ěr AT/GC ............................................................................................................. 31 6. Charakteristika analyzovaných druh ů a vyšších taxon ů ............................................. 33 6.1. Ophioglossales, Ophioglossaceae ..................................................................... 33 6.2. Psilotales , Psilotaceae ...................................................................................... 33 6.3. Equisetales , Equisetaceae ................................................................................. 33 6.4. Marattiales , Marattiaceae ................................................................................. 35 6.5. Osmundales , Osmundaceae .............................................................................. 35 6.6. Gleicheniales , Gleicheniaceae .......................................................................... 36 6.7. Schizaeales , Lygodiaceae .................................................................................. 36 6.8. Schizaeaeles , Anemiaceae ................................................................................. 36 6.9. Salviniales , Marsileaceae ................................................................................. 37 6.10. Salviniales , Salviniaceae ................................................................................... 37 6.11. Cyatheales , Cibotaceae ..................................................................................... 37 6.12. Cyatheales , Cyatheaceae .................................................................................. 38 6.13. Cyatheales , Dicksoniaceae ............................................................................... 38 6.14. Polypodiales , Lindsaeaceae .............................................................................. 38 6.15. Polypodiales , Dennstaedtiaceae ....................................................................... 38 6.16. Polypodiales , Pteridaceae ................................................................................. 38 6.17. Polypodiales , Aspleniaceae ............................................................................... 39 6.18. Polypodiales , Thelypteridaceae ........................................................................ 39 6.19. Polypodiales , Woodsiaceae ............................................................................... 40 6.20. Polypodiales , Blechnaceae ............................................................................... 40 6.21. Polypodiales , Onocleaceae ............................................................................... 40 6.22. Polypodiales , Dryopteridaceae ........................................................................
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