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Univerzita Karlova V Praze Přírodovědecká Fakulta Univerzita Karlova v Praze Přírodovědecká fakulta Studijní program: Biologie Studijní obor: Obecná biologie Diverzita, ekologie a ekofyziologie sněžných řas Diversity, ecology and ecophysiology of snow algae Bakalářská práce Lenka Mikešová Vedoucí práce: RNDr. Linda Nedbalová, Ph.D. Praha, 2013 Prohlášení: Prohlašuji, že jsem závěrečnou práci zpracovala samostatně a že jsem uvedla všechny použité informační zdroje a literaturu. Tato práce ani její podstatná část nebyla předložena k získání jiného nebo stejného akademického titulu. V Praze, 17. 5. 2013 Podpis Poděkování: Na tomto místě bych ráda velmi poděkovala své školitelce RNDr. Lindě Nedbalové, Ph.D. za spoustu cenných rad a trpělivé a přátelské vedení mé bakalářské práce. Poděkování patří i mé rodině za jejich nezbytnou podporu a pomoc během celého studia. Abstrakt Trvalá i dočasná sněhová pokrývka polárních a horských oblastí je velmi extrémním habitatem. Přesto existují organismy, které toto prostředí obývají. Mezi významné zástupce kryosestonu patří zelené řasy z řádu Chlamydomonadales (Chlorophyta), které jsou známé z polárních a horských oblastí celého světa. Tyto organizmy, které obsadily sněžné prostředí pravděpodobně až sekundárně, dokázaly vyvinout různé ekofyziologické adaptace nezbytné k úspěšnému přežití v extrémních podmínkách. Nízké teploty a vysoká intenzita záření jsou hlavními faktory, kterým musí přítomné organizmy čelit. Mezi nejdůležitější mechanismy přežití patří přizpůsobení životních cyklů (například střídání odolných stádií a pohyblivých vegetativních stádií), syntéza a akumulace ochranných sekundárních karotenoidů, změna ve složení mastných kyselin membrán a u některých druhů snížení teplotního optima růstu. Právě tyto adaptace jsou v současnosti spolu se studiem diverzity hlavním předmětem výzkumu sněžných řas. Klíčová slova: Chlamydomonas, Chloromonas, sně žné řasy, živiny, životní cykly, psychrofilní, astaxanthin, polynenasycené mastné kyseliny Abstract Permanent and temporary snow cover in polar and mountain areas is a very extreme habitat. However, there are organisms that inhabit this environment. Green algae of the order Chlamydomonadales (Chlorophyta) belong among the important representatives of cryoseston, which are known from polar and mountain regions around the world. These organisms, which occupied the snow environment probably secondarily, developed different ecophysiological adaptations required for the successful survival in extreme conditions. Low temperatures and high levels of radiation are the main extreme factors that organisms must cope with. The most important survival mechanisms include the adaptation of life cycles (such as of the shifts between resistant stages and moving vegetative stages), accumulation of secondary carotenoids, changes in fatty acid composition of membranes and lowering of growth temperature optima in some species. These adaptations together with the study of diversity represent at present the main subjects of research of snow algae. Key words: Chlamydomonas, Chloromonas, snow algae, nutrients, life cycles, psychrophiles, astaxanthin, polyunsaturated fatty acids Obsah 1. Úvod......................................................................................................................... 2 2. Diverzita................................................................................................................... 3 2.1 Zelené řasy s bičíkatými stádii v životním cyklu (Chlamydomonadales)........ 3 2.2 Vývoj taxonomického postavení sněžných řas z řádu Chlamydomonadales ... 6 2.3 Zelené řasy bez bičíkatých stádií v životním cyklu.......................................... 9 2.4 Ostatní druhy fotoautotrofních mikroorganizmů............................................ 11 2.5 Rozšíření sněžných řas rodů Chlamydomonas a Chloromonas...................... 11 3. Ekologie ................................................................................................................. 13 3.1 Fyzikální vlastnosti sněhu .............................................................................. 13 3.2 Chemické vlastnosti sněhu ............................................................................. 14 3.3 Životní cykly sněžných řas............................................................................. 16 3.4 Interakce s ostatními mikroorganizmy ........................................................... 19 4. Ekofyziologie......................................................................................................... 21 4.1 Teplotní optima růstu...................................................................................... 21 4.2 Sekundární karotenoidy.................................................................................. 23 4.3 Složení mastných kyselin ............................................................................... 27 4.4 Fotosyntéza..................................................................................................... 30 5. Závěr ..................................................................................................................... 34 6. Seznam literatury .................................................................................................. 36 1 1. Úvod Fotoautotrofní mikroorganizmy (prokaryotní i eukaryotní) často najdeme v prostředí s trvale či periodicky se vyskytujícími extrémními hodnotami základních ekologických faktorů, jako je teplota, pH, tlak, salinita či záření (Seckbach 2007). Příkladem habitatu s extrémními podmínkami pro život je sněhová pokrývka, která zaujímá rozsáhlé plochy polárních a horských oblastí. Pro organismy, které nejsou na chlad adaptovány, mohou mít nízké teploty či dokonce teploty pod bodem mrazu za následek řadu letálních poškození buněk. Jedním z nich je zmrazení intracelulární vody, jelikož ledové krystaly mohou roztrhat buněčné struktury. Kvůli absenci vody v kapalném skupenství dochází často k zastavení metabolických procesů (Rothschild a Mancinelli 2001). Ve sněhu dochází ke střídání procesů vymrzání a tání a často bývá velmi chudý na obsah živin (Remias a kol. 2005). Oblasti se sněhovou pokrývkou bývají vystaveny vysoké intenzitě UV a viditelného záření, která může vést k poškození fotosyntetického aparátu a nukleových kyselin (přímo nebo prostřednictvím volných kyslíkových radikálů) (Rothschild a Mancinelli 2001). I přes velmi nehostinné podmínky existují mikroorganizmy, které se na prostředí sněhu specializovaly (Gomes a Steiner 2004). Jedná se především o řasy, které byly nalezeny ve sněhových polích celého světa (Kol 1968). Za tzv. „pravé“ sněžné řasy jsou považovány druhy, které se rozmnožují ve velmi specifickém prostředí, jakým je kapalná voda v tající sněhové pokrývce. Za příznivých podmínek tvoří nápadné barevné sněhy, které jsou významným fenoménem horských a polárních oblastí. Patří mezi ně především zelení bičíkovci z řádu Chlamydomonadales(Chlorophyta) (Stibal a Elster 2005). Sněžné řasy jsou ve svém prostředí významnými primárními producenty, kteří se podílí na koloběhu uhlíku (Williams a kol. 2003). Tyto extremofilní organismy představují unikátní modelovou skupinu pro studium adaptací na nízké teploty. Díky rychlému růstu v nízkých teplotách a produkci cenných organických látek mají také značný biotechnologický potenciál v oblastech enzymologie, molekulární biologie, medicíny, zpracování odpadu či v potravinářství (Hoham a Duval 2001). Bakalářská práce je shrnutím současných znalostí o taxonomickém složení a diverzitě sněžných řas, jejich ekologii a ekofyziologických přizpůsobeních, bez kterých by tyto organizmy nemohly obsadit unikátní habitat sněhové pokrývky. 2 2. Diverzita Barevný sníh byl známý již před mnoha staletími, avšak příčiny tohoto nápadného jevu zůstávaly dlouho neznámé. Nejprve se usuzovalo, že červené zbarvení sněhu je anorganického původu (Kol 1957). V roce 1819 Francis Bauer zkoumal vzorky červeného sněhu z oblasti Baffinova zálivu. Pod mikroskopem pozoroval stovky tmavě červených globulí, které nejprve přirovnával k pylovým zrnům a poté k houbovým organizmům. Pozorovaný organizmus nazval Uredo nivalis (Bauer 1819). Profesor N. Wille jako jeden z prvních vědců pozoroval červený sníh pod mikroskopem přímo na ledovci v Norsku. Zjistil, že původcem nápadného zbarvení jsou mikroorganizmy, konkrétně řasy. Popsal tak celosvětově rozšířený druh Chlamydomonas nivalis (Bauer) Wille, který může tvořit obrovskou biomasu a způsobovat výrazné červené nárosty (Kol 1957). Kol (1968) popsala vegetativní buňky Chlamydomonas nivalis jako elipsovité s centrálním pyrenoidem a dvěma kontraktilními vakuolami. Kulovité cysty mají tenkou zdrsnělou buněčnou stěnu a obsahují velké množství sekundárního karotenoidu astaxanthinu. Na základě těchto cyst s minimem determinačních znaků přítomných v č erveném sněhu bývá druh Chlamydomonas nivalis dodnes ze sněhu popisován. Je proto pravděpodobné, že se nejedná o jeden druh (ačkoliv v současnosti taxonomicky uznávaný), ale o komplex několika různých druhů (viz kapitola 2.2.). Na území bývalého Československa byly cysty Chlamydomonas nivalis poprvé nalezeny roku 1976 v Schustlerově zahrádce v Krkonoších (Fott a kol. 1978). Méně nápadný zelený sníh poprvé pozoroval již v 18. století profesor J.A. Czirbesz v Tatrách. Jako původci tohoto zbarvení bylo popsáno několik vláknitých řas z rodu Raphidonema, např. R. tatrae (Kol) Hind.nebo R. nivale Lagerh., která může způsobovat též růžové zbarvení (Kol 1957). 2.1. Zelené řasy s bičíkatými stádii v životním cyklu (Chlamydomonadales) Řasy tvořící barevné sněhy patří nejčastěji
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