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Diplomová Práce Milan Kotilínek-S

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Diplomová Práce Milan Kotilínek-S Jiho česká univerzita v Českých Bud ějovicích Přírodov ědecká fakulta Pr ůběh klí čení a mykorhizní asociace dvou druh ů rodu Neottia Diplomová práce Bc. Milan Kotilínek Školitel: RNDr. Jana Jersáková, PhD. Konzultant: RNDr. Tamara T ěšitelová České Bud ějovice 2012 Kotilínek, M., 2012. Pr ůběh klí čení a mykorhizní asociace dvou druh ů rodu Neottia . [Germination course and mycorrhizal association of two Neottia species. Mgr. Thesis, in Czech]. – 30p., Faculty of Science, University of South Bohemia, České Bud ějovice, Czech Republic. Anotace Germination course and mycorrhizal partners during germination and adulthood were studied in Neottia ovata and N. cordata . Neottia ovata was studied in two different biotops: meadows and forests. Research was conducted in in situ conditions in the Šumava and the Smr činy mountains. Prohlašuji, že svoji diplomovou práci jsem vypracoval samostatn ě pouze s použitím pramen ů a literatury uvedených v seznamu citované literatury. Prohlašuji, že v souladu s § 47b zákona č. 111/1998 Sb. v platném zn ění souhlasím se zve řejn ěním své diplomové práce, a to v nezkrácené podob ě elektronickou cestou ve ve řejn ě přístupné části databáze STAG provozované Jiho českou univerzitou v Českých Bud ějovicích na jejích internetových stránkách, a to se zachováním mého autorského práva k odevzdanému textu této kvalifika ční práce. Souhlasím dále s tím, aby toutéž elektronickou cestou byly v souladu s uvedeným ustanovením zákona č. 111/1998 Sb. zve řejn ěny posudky školitele a oponent ů práce i záznam o pr ůběhu a výsledku obhajoby kvalifika ční práce. Rovn ěž souhlasím s porovnáním textu mé kvalifika ční práce s databází kvalifika čních prací Theses.cz provozovanou Národním registrem vysokoškolských kvalifika čních prací a systémem na odhalování plagiát ů. V Českých Bud ějovicích dne 14.12.2012 ............................................. II Pod ěkování Na tomto míst ě bych cht ěl pod ěkovat všem, kte ří mi pomáhali p ři studiu mykorhizy a bez jejichž pomoci by tato práce nikdy nevznikla. V první řad ě d ěkuji své školitelce Jan ě Jersákové, která mi umožnila studium orchidejí, za její úžasný p řístup a za rady, které mi usnadnily bádání. Konzultantce Tama ře T ěšitelové děkuji za pomoc s řešením všech možných i nemožných problém ů a bezmeznou trp ělivost b ěhem spolupráce v terénu i v laborato ři. Tama ře také pat ří velký dík za pomoc se statickým vyhodnocením v programu R. Ob ěma také musím pod ěkovat za pomoc p ři sepisování této práce, kdy byly ochotné číst mé výplody a opravovat opakující se gramatické chyby. Také bych cht ěl pod ěkovat Jít ě a Babce za korekci pravopisu, množství rad a jejich po ťouchlé nápady, kterými m ě vždycky pobaví. Také d ěkuji svojí mamin ě za podporu a neustálé zásobování chutnými pokrmy. Dále bych cht ěl pod ěkovat Davidu P ůbalovi z CHKO Šumava a Iv ě Bufkové z NP Šumava za pomoc p ři vyhledávání vhodných lokalit pro výzkum, Bá ře a Françoisovi za pomoc p ři zpracování vzork ů v laborato ři a Jirkovi za dobré rady a ochotu vždy pomoci. Nesmím také zapomenout na naší Škodu 1203 WRC s pohonem na plyn, jež m ě vždy dovezla na lokality i zp ět. A to i p řes své stá ří a ob časný nedostatek pochopení pracovník ů NP Šumava pro ‚veterány‘. V neposlední řad ě také d ěkuji NP a CHKO Šumava za umožn ění výzkumu na jejich území a za ubytování na správ ě NP v Kašperských Horách a GA AV ČR č.IAA600870802 za financování projektu, jehož sou částí je i má diplomová práce. III Obsah 1. Úvod ................................................................................................................................... 1 1.1. Orchideoidní mykorhiza .......................................................................................... 1 1.2. Houboví partne ři ...................................................................................................... 3 1.3. Ekologie klí čení orchidejí ........................................................................................ 4 1.4. Vliv biotopu a genotypu na orchideoidní mykorhizu ........................................... 6 1.5. Ekologie a mykorhizní asociace studovaných druh ů............................................ 6 1.6. Cíle práce .................................................................................................................. 8 2. Metodika ........................................................................................................................... 8 2.1. Popis modelových druh ů ......................................................................................... 8 2.2. Popis lokalit ............................................................................................................... 8 2.3. Vysévací pokus .......................................................................................................... 8 2.4. Analýza symbiotických hub v protokormech ...................................................... 10 2.5. Analýza symbiotických hub v ko řenech dosp ělc ů............................................... 10 2.6. Metody molekulární identifikace .......................................................................... 11 2.7. Statistická analýza dat ........................................................................................... 12 2.8. Fylogenetická analýza dat ..................................................................................... 13 2.9. TEM ......................................................................................................................... 14 3. Výsledky .......................................................................................................................... 15 3.1. Pr ůběh klí čení ......................................................................................................... 15 3.2. Symbiotické houby v protokormech a ko řenech dosp ělc ů................................. 18 3.3. TEM ......................................................................................................................... 24 4. Diskuse ............................................................................................................................. 25 5. Záv ěr................................................................................................................................ 30 6. Literatura ........................................................................................................................ 31 7. Přílohy ............................................................................................................................. 39 8. Fotop říloha ...................................................................................................................... 44 IV 1. Úvod Mykorhizní symbióza je d ůležitou sou částí terestrických ekosystém ů. Udává se, že více než 80 % druh ů rostlin je schopno tvo řit mykorhizu s půdními houbami (Wang a Qiu, 2006). V sou časnosti je rozlišováno n ěkolik typ ů mykorhizy na základ ě zp ůsobu kontaktu mezi ko řeny a houbou. Pro ektomykorhizní symbiózu (ECM) je charakteristický hyfový pláš ť obalující špi čky ko řen ů a tzv. Hartigova sí ť tvo řená houbovými vlákny v mezibun ěč ných prostorech; hyfy však nepronikají do bun ěk. Pokud vlákna pronikají do bun ěk hostitelské rostliny, mluvíme již o endomykorhizní symbióze. Tu m ůžeme dále rozd ělit na arbuskulární mykorhizu, erikoidní mykorhizu a orchideoidní mykorhizu (OM). Pro úplnost je t řeba ješt ě vzpomenout ektendomykorhizu. Ta kombinuje znaky obou p ředchozích a m ůžeme ji dále rozd ělit na mykorhizu arbutoidní, monotropoidní a pyroloidní (Gryndler a kol., 2004). Ze všech výše vyjmenovaných typů mykorhizy se dále budu zabývat převážn ě jen orchideoidní mykorhizou. Běžnou mykorhizu m ůžeme popsat jako mutualistický vztah houby s autotrofní rostlinou, kdy rostlina od houby získává minerální látky a vodu a houb ě p ředává část uhlíku získaného b ěhem fotosyntézy. Existují však výjimky. U p řibližn ě 450 druh ů rostlin náležejících do r ůzných čeledí (Merckx a Freudenstein, 2010) bylo zjišt ěno, že od houby získávají i uhlík a stávají se tak parazity. Tento zp ůsob výživy se nazývá mykoheterotrofie (MH) (Leake, 1994). 1.1. Orchideoidní mykorhiza Tento typ mykorhizy je typický pro všechny druhy z čeledi Orchidaceae, u žádné další čeledi pozorován nebyl (Gryndler a kol., 2004). Čele ď Orchidaceae, s odhadovaným po čtem druh ů 20 000 až 35 000 (Dressler, 1993 a 2004; Malberley, 1997; Cribb a kol., 2003); je v sou časné dob ě považována za nejv ětší čele ď v ůbec (Cribb a kol., 2003; WCSP, 2012). Vývojov ě se jedná o izolovanou v ětev řádu Asparagales, která se odd ělila p řed 76 až 84 mil. let (Ramírez a kol., 2007). Orchideje mají centrum rozší ření v tropech, ale m ůžeme je nalézt po celé zem ěkouli až po boreální pásmo. M ůžeme najít jak terestrické, tak i epifytické a litofytické r ůstové formy. Navzdory této rozmanitosti spojuje všechny orchideje jejich závislost na mykorhizních houbách b ěhem klí čení, kdy semena nejsou schopna klí čit bez přítomnosti vhodné symbiotické houby (nap ř. Rasmussen, 1995; Gryndler a kol., 2004; Smith a Read, 2008). Další v ěcí, unikátní pro všechny orchideje, je útvar nazývaný protokorm. Jedná se o specifický klí ční útvar orchidejí a odpovídá hypokotylu a ko řenovému základu u ostatních rostlin. Vzniká z embrya a pozd ěji se diferencuje v jednotlivá pletiva. 1 Orchideoidní mykorhiza byla poprvé popsána na po čátku 20. století Bernardem (1909) a Burgeffem (1909). Jimi popsaná forma je nazývána tolypofágní. Pozd ěji pak byla ješt ě popsána daleko
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