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Bakalářská Práce MASARYKOVA UNIVERZITA PŘÍRODOVĚDECKÁ FAKULTA ÚSTAV BOTANIKY A ZOOLOGIE Bakalářská práce Brno 2012 Ondřej Knápek MASARYKOVA UNIVERZITA PŘÍRODOVĚDECKÁ FAKULTA ÚSTAV BOTANIKY A ZOOLOGIE Evoluce velikosti genomu a obsahu DNA bazí v rodě Nepenthes Bakalářská práce Ondřej Knápek Vedoucí práce: Mgr. Petr Šmarda, PhD. Brno 2012 Bibliografický záznam Autor: Ondřej Knápek Přírodovědecká fakulta, Masarykova univerzita Ústav botaniky a zoologie Evoluce velikosti genomu a obsahu DNA bazí v rodě Název práce: Nepenthes Studijní program: PřF B-BI Biologie Studijní obor: PřF BISE Systematická biologie a ekologie Vedoucí práce: Mgr. Petr Šmarda, Ph.D. Akademický rok: 2011/2012 Počet stran: 41+7 Klíčová slova: průtoková cytometrie, láčkovka, Nepenthes, velikost genomu, zastoupení AT/GC bazí, délka průduchů Bibliographic Entry Author: Ondřej Knápek Faculty of Science, Masaryk University Department of Botany and Zoology Evolution of genome size and DNA base composition in Title of Thesis: Nepenthes Degree programme: PřF B-BI Biology Field of Study: PřF BISE Systematic Biology and Ecology Supervisor: Mgr. Petr Šmarda, Ph.D. Academic Year: 2011/2012 Number of Pages: 41+7 Keyword: flow cytometry, pitcher plant, Nepenthes, genome size AT/GC content, stomatal length Abstrakt Rod Nepenthes je tropickým rodem masožravých rostlin s cca 120 druhy, rostoucími v pralesních a horských oblastech převážně v Indomalajsii. Pestrost stanovišť, ekologie, geografická izolace ostrovů a masožravá strategie výživy, zde umožňuje testovat řadu hypotéz souvisejících s evolucí genomu rostlin za různých podmínek prostředí. Velikost genomu a AT/GC genomový poměr byly změřeny pro 109 vzorků 97 druhů rodu Nepenthes. Z rostlin byly odebrány vzorky mladých listů pro cytometrická měření a použitím mikroreliéfové metody byly také sejmuty u starších listů otisky spodních stran listů pro měření průduchů. K analyzovaným druhům byly z dostupné literatury dohledány informace o ekologii, morfologii a rozšíření. Velikost genomu u Nepenthes se pohybuje od 1,37 5,95 pg a obsah GC bazí je v rozsahu 35,54 45,13 %, což je v případě velikosti genomu nejvíce u zatím známých masožravých rostlin. Menší velikosti genomu mají primitivní druhy na západním okraji areálu rodu a nejvetší pak radiující druhy, zejména na Sumatře a Borneu, kde by některé druhy (N. lavicola, N. bicalcarata) mohly být polyploidní. Větší velikost genomu na ostrovech naznačuje, že místní radiace by mohla být spojena s amplifikací GC bohaté DNA, vzhledem k signifikantní korelaci velikosti genomu a obsahu GC. Velikost průduchů v rodu je ovlivněna jak velikostí genomu, tak prostředím ve kterém se druhy vyskytují. Abstract Nepenthes is a tropical genus of carnivorous plants comprising about 120 species, growing in rainforest and montane areas of Indomalaysia. The diversity of habitats, ecology, geographical isolation of islands and carnivorous nutrition strategy, allows testing a number of hypotheses related to the evolution of plant genome under various environmental conditions. Genome size and AT / GC genomic ratio was measured for 109 samples of 97 species of the genus Nepenthes. Samples of young leaves were collected for cytometric measurements. Microrelief reprint method was used for the measurement of the leaf stomatal length. Data about ecology, morphology and distribution were obtained from the available literature. Genome size of Nepenthes ranges from 1.37 up to 5.95 pg; the content of GC bases ranges from 35.54 to 45.13%, which is the largest genome size yet known in carnivorous plants. Smaller genome size are found in the primitive species in the western edge of Nepenthes distribution area and larger genome sizes are typical of radiating species, in particular on Sumatra and Borneo island, where some species with large genomes (N. lavicola, N. bicalcarata) could be polyploids. Larger genome size on the islands suggests that local radiation may be associated with amplification of GC-rich DNA, as supported by the significant correlation of genome size and GC content. Stomatal size in the genus seems affected both by genome size and habitat conditions. Poděkování Na tomto místě bych chtěl poděkovat všem, kteří se fyzicky či psychicky podíleli na zdárném dokončení této práce. Na prvním místě bych tedy chtěl poděkovat mému vedoucímu, panu Mgr. Petru Šmardovi, Ph.D.1) za ochotu při konzultacích a trpělivost při vysvětlování; dále pak patří velký dík Mgr. Lucii Horové1) a Ing. Alexandře Březinové1) za pomoc s cytometrickými měřeními a Mgr. Adamu Velebovi1) za fotografie druhů. Také bych chtěl moc poděkovat RNDr. Vlastiku Rybkovi Ph.D.2) a Mgr. Martinu Dančákovi, Ph.D.3) za poskytnutí rostlinného materiálu k měřením a pomoc při dotváření výsledné matice znaků. Bez jejich ochotné spolupráce by tato studie nevznikla. Děkuji také všem, kteří mi dodávali dobrou náladu a optimismus. Finanční zázemí této práce zajistily projekty LC06073 a MSM0021622416 poskytnuté Ministerstvem školství, mládeže a tělovýchovy a projekt GAČR505/11/0881 poskytovaný Grantovou agenturou České republiky. 1) Ústav botaniky a zoologie, Přírodovědecká fakulta Masarykovy univerzity v Brně 2) Botanická zahrada Praha–Trója 3) Katedra botaniky, Přírodovědecká fakulta Univerzity Palackého v Olomouci Prohlášení Prohlašuji, že jsem svoji bakalářskou práci vypracoval samostatně s využitím informačních zdrojů, které jsou v práci citovány. Souhlasím s uložením této bakalářské práce v knihovně Ústavu botaniky a zoologie PřF MU v Brně, případně v jiné knihovně MU, s jejím veřejným půjčováním a využitím pro vědecké, vzdělávací nebo jiné veřejně prospěšné účely, a to za předpokladu, že převzaté informace budou řádně citovány a nebudou využívány komerčně. Brno 9. května 2012 ……………………………… Jméno Příjmení Obsah 1. Čeleď Nepenthaceae . 10 1.1. Taxonomie . 10 1.2. Historie výzkumu . 10 1.3. Morfologie . 11 1.4. Rozšíření . 12 1.4.1. Teorie osidlování indomalajské oblasti . 13 1.5. Ekologie . 14 1.6. Pasti (láčky) . 15 2. Obsah DNA a poměr AT/CG bazí. 18 2.1. Velikost genomu . 18 2.2. Poměr AT/CG bazí . 19 2.3. Velikost průduchů . 20 3. Cíle práce . 21 4. Materiál a metodika . 22 4.1. Materiál . 22 4.2. Průtoková cytometrie . 22 4.3. Mikroreliéfová metoda . 23 4.4. Newick formát, TreeGraph, Phylocom. 23 4.5. Statistická analýza . 25 5. Výsledky . 26 6. Diskuse . 32 7. Literatura . 34 8. Přílohy . 41 - 9 - 1. Čeleď Nepenthaceae 1.1. Taxonomie Čeleď Nepenthaceae Dumortier patří do řádu Caryophyllales Berchtold & J. Presl a obsahuje pouze jediný rod Nepenthes - láčkovka čítající přibližně 120 druhů (McPherson 2009). Kromě toho je známa i řada přírodních a umělých hybridů. Typovým druhem rodu je Nepenthes distillatoria L. 1.2. Historie výzkumu Rostliny z rodu Nepenthes poprvé zmiňuje Etienne de Flacourt (1658), francouzský guvernér na Madagaskaru. Láčkotvaré rostlině popisované v jeho práci Histoire de la Grande Isle de Madagascar dal jméno Amramatico, pocházející z jazyka místních domorodců. Validně byla rostlina popsána až r. 1797 Jeanem Louisem Maria Poiretem jako N. madagascariensis (Poiret 1797). Další zmínku o rostlině z tohoto rodu, tentokrát ze Srí Lanky, najdeme v práci Acta Medica et Philosophica Hafniensia (Thomas Bartolin 1677; pod jménem „miranda herba“ = „podivuhodná bylina“). Validně tento druh pak popisuje Carl von Linné (1753) pod jménem Nepenthes distillatoria L. Roku 1790 popsal portugalský kněz João Loureiro, ve svém díle Flora Cochinchinensis, druh Phyllamphora mirabilis de Loureiro (1790), dnes známý jako N. mirabilis (Druce 1916). Mnoho druhů láčkovek bylo objeveno a popsáno zejména během řady botanických výprav začátkem 19. století (McPherson 2009). Množství druhů těchto podivuhodných rostlin upoutalo pozornost nejen přírodovědců, ale také veřejnosti, hlavně v tehdejší viktoriánské Anglii, kde pokusy o pěstování láčkovek sahají do roku 1789 (Veitch 1897). Zájem veřejnosti vyvolal také snahu pěstovat láčkovky komerčně. Nejznámější pěstírnou byla bezpochyby firma Veitch Nurseries. Ta podnikala objevné expedice a zabývala se sběrem a dovozem semen i celých rostlin do Velké Británie. Zde je pěstovala ve velkých vytápěných sklenících a dodávala je zprvu botanickým institucím, později pak i bohatým nadšencům a nakonec i širší veřejnosti. V polovině 19. století již láčkovky byly celkem běžně pěstovány v botanických zahradách Velké Británie. První monografii rodu, čítající tehdy 22 druhů, sepsal Ernst Wunschmann (1872). Tuto monografii posléze doplnil o dalších 11 druhů Sir Joseph - 10 - Dalton Hooker. Jeho práce Nepenthaceae, zaměřená hlavně na druhy z Bornea, byla publikována v díle Prodromus systematis universalis regni vegetabilis (Hooker in de Candolle 1873). O další monografické zpracování rodu se zasloužil John Muirhead Macfarlane (1908). Jeho práce tehdy popisovala 58 druhů a navíc všechny tehdy známé umělé hybridy. V pozdější monografii Benedictus Danser (1928) sjednocením taxonomickéhho pojetí a synonymiky zredukoval původní počet Macfarlanových 58 druhů na 47, zároveň také přidal popis dalších 17 nových druhů, pocházejících hlavně z Indonésie, Malajsie a Nové Guiney. Ze Sumatry popsali 18 nových druhů Tamin a Hotta (1986), i když se ale později ukázalo, že některé z těchto druhů představují pouze přírodní hybridy. K novějším monografiím rodu patří práce Matthew Jebb & Martin Cheek (1997). Australský botanik a ekolog Charles Clarke zkoumal láčkovky ze Sumatry, Bornea a Malajského poloostrova. Své poznatky publikoval ve dvou knihách Nepenthes of Borneo (Clarke 1997) a Nepenthes of Sumatra and Peninsular Malaysia (Clarke 2001). Poslední ucelený přehled
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