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Přírodovědecká Fakulta Je Rozšíření Orchidejí Limitováno Jejich

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Přírodovědecká Fakulta Je Rozšíření Orchidejí Limitováno Jejich Jiho česká univerzita v Českých Bud ějovicích Přírodov ědecká fakulta Bakalá řská práce Je rozší ření orchidejí limitováno jejich mykorhizními partnery? Milan Kotilínek Školitel: RNDr. Jana Jersáková, PhD. České Bud ějovice 2010 Kotilínek M. 2010. Je rozší ření orchidejí limitováno rozší řením jejich mykorhizních partner ů? [Is orchid distribution limited by their mycorrhizal associations? Bc. Thesis, in Czech]. – 25p., Faculty of Science, The University of South Bohemia, České Bud ějovice, Czech Republic. Anotace The first part of this thesis is a review on limitations of orchids by their mycorrhizal symbiosis. The second part presents on outline of my master thesis, which is focused on ecology and mycorrhizal partners of two Listera species. Prohlašuji, že svoji bakalá řskou 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 bakalá řské 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. V Českých Bud ějovicích dne 30. dubna 2010. .................................................... II Pod ěkování Na tomto míst ě bych cht ěl pod ěkovat všem, kte ří mi b ěhem posledních t ří let pomáhali při studiu mykorhizy. V první řad ě děkuji své školitelce Jan ě Jersákové za uvedení do taj ů mykorhizní symbiózy u orchidejí a množství rad, které mi usnadnily bádání; a Tama ře Malinové za pomoc s řešením všech možných i nemožných problém ů a bezmeznou trp ělivost během spolupráce v terénu i v laborato ři. Velký dík také pat ří ob ěma za pomoc p ři sepisování této práce. Dále bych cht ěl poděkovat 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 mojí mamince za vzdálenou podporu skrze chutné zásoby jídla. Děkuji také Pet ře za psychickou podporu b ěhem psaní a Ond řejovi za gramatickou korekci tohoto textu. 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 část je sou částí mé diplomové práce. III Obsah 1. Úvod ................................................................................................................ - 1 - 1.1 Mykorhizní symbióza ................................................................................ - 1 - 2. Orchideoidní mykorhiza ................................................................................... - 3 - 2.1 Základní popis OM ................................................................................... - 3 - 2.2 Význam mykorhizy pro orchideje .............................................................. - 4 - 2.3 Houboví partne ři ....................................................................................... - 6 - 3. Fyziologie OM ................................................................................................. - 7 - 3.1 Přehled stadií klí čení a jejich závislosti na mykorhize .............................. - 7 - 3.2 Přenos uhlíku mezi houbou a rostlinou .................................................... - 8 - 3.3 Přenos dusíku a dalších látek mezi houbou a rostlinou ............................ - 9 - 4. Abiotické a biotické faktory ovliv ňující klí čení a r ůst orchidejí ........................ - 10 - 4.1 Abiotické faktory ..................................................................................... - 10 - 4.2 Biotické faktory ....................................................................................... - 11 - 5. Ekologické d ůsledky závislosti na mykorhizní symbióze ............................... - 12 - 6. Záv ěr ............................................................................................................. - 14 - 7. Návrh diplomové práce .................................................................................. - 15 - 7.1 Úvod ....................................................................................................... - 15 - 7.2 Metodika ................................................................................................. - 15 - 7.2.1 Popis vybraných druh ů: ................................................................... - 15 - 7.2.2 Popis lokalit ..................................................................................... - 16 - 7.2.3 Vysévací pokus ............................................................................... - 16 - 7.2.4 Mykorhizní partne ři v dosp ělosti ...................................................... - 18 - 7.2.5 Metody molekulární identifikace ...................................................... - 18 - 7.2.6 Izotopová analýza ........................................................................... - 19 - 7.3 Výsledky ................................................................................................. - 20 - 7.3.1 Pr ůběh klí čení ................................................................................. - 20 - 7.3.2 Mykorhizní partne ři v ko řenech dosp ělých rostlin ............................ - 20 - 7.3.3 Izotopová analýza ........................................................................... - 20 - 7.4 Diskuse .................................................................................................. - 23 - 8. Literatura ....................................................................................................... - 25 - Přílohy .................................................................................................................. - 31 - IV 1. Úvod 1.1 Mykorhizní symbióza Mykorhizní symbióza je druh soužití mezi houbou a rostlinou, p ři kterém vznikají specifické orgány zvané mykorhizy. Mykorhizní symbióza je v přírod ě široce rozší řena a odhaduje se, že asi 90 % všech rostlin je běžn ě spojeno s půdními houbami (Wang a Qiu 2006). Jedná se tak o důležitou sou část terestrických ekosystémů. V klasickém pojetí mykorhizní symbiózy dodává houba rostlin ě minerální látky a vodu, rostlina naopak poskytuje houb ě asimiláty. Jedná se tak o vztah mutualistický. Houby, vyskytující se v mykorhizách, zahrnují n ěkolik skupin pravých hub (Eumycota), a to Ascomycota, Basidiomycota, a Glomeromycota (Smith a Read 2008). Tyto houby kolonizují hlavn ě ko řenovou pokožku (rhizodermis) a ko řenovou k ůru, která zpravidla tvo ří několik vrstev pod rhizodermis (Gryndler a kol. 2004). Mykorhizy rozd ělujeme podle zp ůsobu kontaktu mezi houbou a ko řenem. U endomykorhizního typu pronikají houbová vlákna do bun ěk hostitelské rostliny, u ektomykorhizy se houbová vlákna nacházejí jen v mezibun ěč ných prostorech. Pokud mykorhiza kombinuje znaky obou výše uvedených typ ů, nazýváme ji ektendomykorhiza (Gryndler a kol. 2004). Endomykorhiza se dále rozd ěluje na n ěkolik dalších typ ů (Gryndler a kol. 2004): • Arbuskulární mykorhiza (AM) je v přírod ě nejrozší řen ější, vyskytuje se přibližn ě u 80% rostlin (Smith a Read 2008) zahrnujících krytosemenné a nahosemenné rostliny, kapradiny, plavun ě a p řesli čky. AM tvo ří houby ze skupiny Glomeromycota. Tato mykorhiza je charakterizována mezibun ěč nými i vnitrobun ěč nými hyfami a ke říčkovitými útvary nazývanými arbuskuly, nacházejícími se uvnit ř bun ěk ko řene. • Erikoidní mykorhiza (ER) se vyskytuje hlavn ě u řádu Ericales a vyzna čuje se charakteristickými smotky hyf uvnit ř bun ěk. Nejvíce jsou p ři ní kolonizovány bu ňky rhizodermis. • Orchideoidní mykorhiza (OM) se vyskytuje u rostlin z čeledi a je charakterizována Orchidaceae vytvá řením klubí ček hyf (peloton ů) uvnit ř bun ěk ko řenové k ůry, skrz rhizodermis houbová vlákna jen pronikají (najdou se ale i výjimky u n ěkterých tropických druh ů (Martos a kol. 2009) u kterých jsou kolonizovány i bu ňky rhizodermis). - 1 - Ektomykorhiza (ECM) je charakterizována hyfovým plášt ěm obalujícím zkrácené ko řínky rostlin a Hartigovou sítí tvo řenou houbovými vlákny v mezibun ěč ných prostorech; vlákna zárove ň nepronikají do vnit řních prostor ů bun ěk hostitele. Tato mykorhiza se vyskytuje tém ěř výlu čně u d řevin (Gryndler a kol. 2004). Mezi mén ě známé druhy mykorhizy pat ří arbutoidní, monotropoidní a pyroloidní mykorhiza. P ři arbutoidní mykorhize obaluje houba ko řeny rostliny hyfovým plášt ěm a vniká do bun ěk rhizodermis. Tuto mykorhizu m ůžeme pozorovat u n ěkterých rostlin z čeledi ∗ Ericaceae. Další dva typy, monotropoidní ( čele ď Monotropaceae ) a pyroloidní ( čele ď ∗ Pyrolaceae ) mykorhiza, jsou si velice podobné. Ko řen je p ři nich obalován hyfovým plášt ěm a v mezibun ěč ných prostorech je vytvá řena Hartigova sí ť, podobn ě jako p ři ECM. Do vnit řních prostor ů bun ěk zasahují hyfy jen krátkými výr ůstky nazývanými kolí čky (Gryndler a kol. 2004). Ve všech t řech p řípadech se jedná o ektendomykorhizy. Na mykorhizní symbiózu však nelze nahlížet jako na ideální mutualismus, spíše se jedná o kontinuum mutualistických až parazitických interakcí, v závislosti na druhu houby, rostliny a podmínkách prost ředí. Dále je mnoho rostlin známo svojí závislostí na houbové výživ ě b ěhem klí čení a po čáte čního vývoje, pozd ěji se však stávají autotrofními. Mezi tyto tzv. po čáte ční mykoheterotrofy řadíme některé kapradiny (nap ř. Botrychium , Ophioglossum ) a plavun ě (nap ř. Lycopodium , Huperzia ), všechny autotrofní druhy orchidejí
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