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Lenka Šafářová POLYPLOIDNÍ KOMPLEX ALLIUM OLERACEUM L

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Lenka Šafářová POLYPLOIDNÍ KOMPLEX ALLIUM OLERACEUM L UNIVERZITA PALACKÉHO V OLOMOUCI PŘÍRODOV ĚDECKÁ FAKULTA KATEDRA BOTANIKY Lenka Šafá řová POLYPLOIDNÍ KOMPLEX ALLIUM OLERACEUM L. V EVROP Ě POLYPLOID COMPLEX OF ALLIUM OLERACEUM L. IN EUROPE DOKTORSKÁ DISERTA ČNÍ PRÁCE Studijní obor botanika Školitel: RNDr. Martin Duchoslav, Ph.D. Olomouc 2011 Bibliografická identifikace Jméno a p říjmení autora: Lenka Šafá řová Název práce: Polyploidní komplex Allium oleraceum L. v Evrop ě Typ práce: doktorská Pracovišt ě: Katedra botaniky – UP Olomouc, Východo české muzeum v Pardubicích Studijní program: Biologie Studijní obor: Botanika Školitel: RNDr. Martin Duchoslav, PhD. Rok obhajoby: 2011 Abstrakt: Polyploidie je p řírodní proces, hrající významnou úlohu v evoluci rostlin, většina z nich ve své minulosti pravd ěpodobn ě prod ělala genovou duplikaci následovanou procesem diploidizace. S využitím metody pr ůtokové cytometrie významn ě stoupla možnost studia polyploidie, struktury polyploidních komplex ů i cytotypové variability jednotlivých rostlinných druh ů. Jako modelový organizmus pro zkoumání polyploidie byl zvolen česnek planý (Allium oleraceum L). Jedná se o polyploidní komplex, u n ěhož byly na po čátku výzkumu známé čty ři ploidní úrovn ě: triploidní (2n=3x=24), tetraploidní (2n=4x=32), pentaploidní (2n=5x=40) a hexaploidní (2n=6x=48). Sb ěr vzork ů probíhal na t řech odlišných prostorových škálách – na úrovni populací, na úrovní p řibližného st ředu rozší ření druhu (Česká Republika, Slovensko) a na úrovni Evropy. Celkem bylo studováno 620 populací a metodou pr ůtokové cytometrie analyzováno 7354 rostlin. Byly potvrzeny všechny doposud známé ploidní úrovn ě, nov ě byly zjišt ěny heptaploidní populace (2n=7x=56) a jedenkrát byla nalezena populace oktoploidní (2n=8x=64). Na úrovni celého areálu druhu jsou dominantním cytotypem pentaploidi následovány tetraploidy a hexaploidy. Triploidní a heptaploidní cytotypy jsou extrémn ě vzácné. Významn ě p řevažují populace pouze s jedním cytotypem, daleko menší množství je populací se dv ěma a se t řemi cytotypy. Populace s více jak t řemi cytotypy nebyly nalezeny. U jednotlivých ploidních úrovní byla zjišt ěna ekologická i prostorová diferenciace. Nejmarkantn ější rozdíl mezi populacemi r ůzných ploidních úrovní je ve stupni ovlivn ění habitat ů člov ěkem, se vr ůstajícím stupn ěm ploidie nar ůstá i stupe ň ruderalizace stanoviš ť. Ploidní úrovn ě se rovn ěž liší v klimatických charakteristikách jako je pr ůměrná ro ční teplota a pr ůměrné ro ční množství srážek. Výše uvedená fakta mohou být d ůkazy pro tzv. adaptivní model diferenciace mezi jednotlivými ploidními úrovn ěmi. Vzhledem k relativn ě četné existenci smíšených populací je však t řeba brát v úvahu i další faktory ovliv ňující výskyt jednotlivých ploidií. Vliv na sou časné rozší ření má patrn ě vysoký stupe ň asexuální reprodukce a omezená schopnost ší ření na v ětší vzdálenosti, odlišné využívání krajiny v minulosti a v neposlední řad ě opakovaný vznik polyploid ů v r ůzných částech areálu. U vybraných jedinc ů všech ploidních úrovní byl zm ěř en absolutní obsah DNA. Monoploidní obsah DNA signifikantn ě klesal se vzr ůstající ploidní úrovní (krom ě heptaploid ů, jejichž monoploidní obsah DNA se nelišil od hexaploid ů). U tetra- a hexaploidního cytotypu byla zjišt ěna variabilita ve velikosti jaderného genomu, která byla těsn ě spjata se zem ěpisnou ší řkou a délkou. Klí čová slova: Allium oleraceum , polyploidie, ploidní úrove ň, cytotyp Po čet stran: 117 Jazyk: český, anglický 2 Bibliographic identification Author’s first name and surname: Lenka Šafá řová Title of the Thesis: Polyploid complex of Allium oleraceum L. in Europe Type of Thesis: Ph.D. Thesis Department: Department of Botany – UP Olomouc, East Bohemian Museum in Pardubice Study program: Biology Field of Study: Botany Supervisor: RNDr. Martin Duchoslav, Ph.D. Year of defence: 2011 Abstract: Polyploidy is a natural process that play a major role in the evolution of majority groups of plants. Most plants have probably undergone past polyploidization followed by diploidization. With use of flow cytometry techniques strongly increase the possibilities for studying polyploid, polyploid complexes or cytotype variation of single species. We choose field garlic ( Allium oleraceum L.) as a model plant for study polyploidy. Allium oleraceum is polyploid complex comprised four documented ploidy levels: triploid (2n=3x=24), tetraploid (2n=4x=32), pentaploid (2n=5x=40) and hexaploid (2n=6x=48). Research run on three different spatial scale: on the level of the population, on the level of the approximate center of the distribution (Czech Republic, Slovak Republic) and on whole natural area distribution (Europe). We sampled 620 populations and by flowcytometry analyzed 7354 plants. We confirmed all of the known ploidy levels. In addition, newly were found heptaploids (2n=7x=56) and once were detected octoploids (2n=8x=64). On the level of whole distribution area dominated pentaploids, followed by tetraploids and hexaploids. Triploids and heptaploids were extremely rare. Significantly predominated single cytotype population, much more less were found two and free cytotype per population. Never were found population with more then three cytotypes. Single cytotypes differ ecologically as spatially. Research revealed considerable variation in niche breath and optimum of cytotypes. Main differentiation among the cytotypes was related to the level of anthropic pressure. The ruderal character of cytotypes increase with a ploidy level. Cytotypes differ also in climatic characteristic like annual mean temperature or annual precipitation. These results could be evidence for adaptive differences among ploidy levels, which may influence their distribution pattern. Local coexistence of cytotypes may support the prevalence of asexual reproduction connected with limited dispersal, equilibrium disrupting processes and repeated origin of the polyploids (unreduced gametes). Selected individuals for single ploidy level were subjected genome size measurement. The monoploid DNA amount (1Cx) was significantly dependent on ploidy level and genome downsizing between two successive cytotypes was significant, except for nonsignificant differences between hexa- and heptaploids. In tetraploids and pentaploids we found intra-ploidy variation in 2C DNA, these variation were tightly connected with longitude and latitude. Key-words: Allium oleraceum , polyploidy, ploidy level, cytotype Number of pages: 117 Language: Czech, English 3 Prohlášení Prohlašuji, že jsem tuto práci vypracovala osobn ě spolu s uvedenými spoluautory za použití citované literatury. Vymezení podílu jednotlivých spoluautor ů 1. Vědecký článek: Complex distribution patterns, ecology and coexistence of ploidy levels of Allium oleraceum (Alliaceae) in the Czech Republic. – Annals of Botany. M. Duchoslav a L. Šafá řová realizovali sb ěr dat a jejich statistické zpracování. F. Krahulec pomohl se sb ěrem části populací. L. Šafá řová provedla laboratorní analýzy. Na tvorb ě textu se podíleli všichni uvedení auto ři. 2. Vědecký článek: Cytotype distribution in mixed populations of polyploid Allium oleraceum measured at a microgeographic scale. – Preslia. L. Šafá řová a M. Duchoslav provedli sb ěr dat. L. Šafá řová provedla laboratorní analýzy a statistické výpo čty, oba auto ři se podíleli na tvorb ě textu. 3. Vědecký článek: Allium oleraceum in Slovakia: cytotype distribution and ecology – Preslia. Všichni auto ři se podíleli na sb ěru dat. L. Šafá řová a M. Jandová provedly laboratorní analýzy, M. Duchoslav provedl statistickou analýzu dat, F. Krahulec vytvo řil první verzi textu, která byla L. Šafá řovou a M. Duchoslavem dopracována. 4. Manuskript: Large cytotype diversity, contrasting distributional patterns and genome size variations in polyploid geophyte Allium oleraceum (Alliaceae) on European scale. L. Šafá řová a M. Duchoslav provedli sb ěr v ětší části populací, L. Šafá řová provedla laboratorní analýzy, v ětší část statistické analýzy dat a oba auto ři se podíleli na tvorb ě textu. Jednotliví spoluauto ři byli seznámeni s výše uvedeným textem a vymezením podílu na jednotlivých článcích. Se zn ěním prohlášení souhlasí. 4 Pod ěkování Na prvním míst ě bych ráda pod ěkovala svému školiteli Martinu Duchoslavovi za to, že mi umožnil spolupracovat na projektu zabývajícím se polyploidním komplexem Allium oleraceum . Velký dík mu pat ří za všechen jeho čas a cenné rady, které mi v ěnoval. Pod ěkování náleží prof. Františku Krahulcovi za jeho nemalou spolupráci p ři sb ěru česnek ů a p ředevším za myšlenkový p řínos, který pro projekt p ředstavuje. Pod ěkování za spolupráci a p řátelství pat ří i koleg ům z řešitelského týmu, především Martin ě Fialové, Michaele Jandové, Alence Jírové, Han ě Sta ňkové a Ji římu Ohryzkovi. Zásadní pro tuto práci bylo technické zázemí poskytnuté laborato ří pr ůtokové cytometrie na Kated ře botaniky Univerzity Palackého v Olomouci – Holici. D ěkuji prof. Aleši Lebedovi za možnost práce v této laborato ři a dr. Iv ě Doleželové za ob ětavou pomoc při mnoha t ěžkých chvílích u pr ůtokového cytometru. V neposlední řad ě bych ráda pod ěkovala všem sb ěratel ům a p řisp ěvatel ům do kolekce evropských česnek ů – p ředevším pak Martinu Dan čákovi, Bohumilu Trávní čkovi, Zby ňkovi Hradílkovi a Janu Mládkovi, jejich úplný seznam lze nalézt v elektronických apendixech jednotlivých článk ů. Výzkum byl podpo řen t ěmito granty: GA ČR, 206/01/P097 GA ČR, 206/04/P115 GA ČR, 206/09/1126 Interní grant UP, PřF 2010/1, 2011/3 5 Obsah 1. ÚVOD............................................................................................................................................................7
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