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Diplomová Práce Přírodovědecká fakulta Masarykovy univerzity Ústav experimentální biologie Oddělení fyziologie a anatomie rostlin DIPLOMOVÁ PRÁCE Efektivní koncentrace polyolů v mechanismech odolnosti lišejníků vůči nízkým teplotám Lubomír Smejkal Vedoucí diplomové práce: Mgr. Josef Hájek, Ph.D. Konzultant: Prof. Ing. Miloš Barták, CSc. Brno 2010 1 Motto: Podle jedné legendy vznikly lišejníky takto: Kojot se pokoušel ulovit labuť. Ta však vzlétla a kojota vzala s sebou do vzduchu. Upustila ho až ve velké výšce a kojot cestou k zemi padal skrz hustý les. A jak se jeho tělo brzdilo o jednotlivé větve, tak se do nich jeho srst stále víc zamotávala. Potom proměnil kojot své chlupy v lišejník slovy: „Má srst nesmí být zmařena. Lidé tě budou sbírat a připravovat z tebe jídlo“. [Turner N. J., Bouchard R., Kennedy D. I. D. (1980): Ethnobotany of the Okanagan-Colville Indians of British Columbia and Washington. Occasional Papers of the British Provincial Museum, Nb. 21: 1–179] 2 Poděkování: Moje poděkování patří především: Mgr. Josefu Hájkovi, Ph.D. (vedoucí) za odborné vedení a cenné rady v průběhu mého magisterského studia a připomínek při zpracování diplomové práce Prof. Ing. Miloši Bartákovi, CSc. (konzultant) za pomoc a jeho odborné rady při experimentální práci v průběhu magisterského studia a při zpracování diplomové práce Mgr. Peterovi Váczimu, Ph.D. za jeho neocenitelné rady při experimentální práci v průběhu magisterského studia RNDr. Stanislavu Manovi, Ph.D. za jeho pomoc při ověřování správnosti chemických vzorců polyolů Doc. RNDr. Marii Kummerové, CSc. za její neocenitelné rady a odborné vedení při nacvičování prezentační činnosti své diplomové a experimentální práce Všem ostatním zaměstnancům Katedry fyziologie a anatomie rostlin PřF MU. V neposlední řadě bych rád poděkoval svojí rodině a přítelkyni za jejich svatou trpělivost a neocenitelnou podporu v průběhu celého studia. 3 Prohlášení Prohlašuji, že jsem zadanou diplomovou práci na téma: „Efektivní koncentrace polyolů v mechanismech odolnosti lišejníků vůči nízkým teplotám” zpracoval samostatně za použití citované literatury. Brno 2010 ________________________ Lubomír Smejkal 4 Abstrakt V mé bakalářské práci jsem se zabýval otázkami souvisejícími s polyoly (cukerné alkoholy), respektive jejich rolí v ochraně lišejníků za nízkých a mrazových teplot. Součástí této práce byl experiment zaměřený na vliv různé koncentrace ribitolu na lišejníky. Základním cílem diplomové práce bylo odpovědět na otázku, jaká je efektivní koncentrace polyolů v lišejníkových stélkách v přirozeném prostředí při působení nízkých teplot. A to je hlavním bodem, kterým bych se měl v mé diplomové práci zabývat. Koncem listopadu 2006 jsme u hráze na Brněnské přehradě sesbírali vzorky lišejníku Xanthoparmelia somloensis (Gyelnik) Hale. Stélky jsme očistili a následně nechali vysychat při pokojové teplotě. Po vyschnutí jsme u poloviny stélek provedli extrakci sekundárních metabolitů, včetně polyolů, pomocí 100% acetonu (varianta aceton rinsing = AR). Druhou polovinu stélek jsme nepromývali (varianta non–AR). Poté jsme stélky zvážili a následně je exponovali po dobu 10 minut v příslušných koncentracích ribitolu, tj. 32mM (5 g/l), resp. 50mM (7,8 g/l). Kontrolní stélky byly po stejnou dobu vlhčeny pouze v destilované vodě. Všechny exponované stélky jsme následně opět zvážili. Tento postup se po 24 hodinách opakoval. Následně jsme stélky umístili do Petriho misek, uzavřeli a oblepili parafilmem (pro zabránění vysychání). Poté jsme Petriho misky umístili do kultivátoru s řízenou teplotou LABIO (CZ) a exponovali za mírné ozářenosti (300 mmol.m-2.s-1) za teplot –5, 0, +5 °C po dobu 8 dní. V průběhu experimentu byly misky několikrát krátkodobě otevřeny a stélky roseny destilovanou vodou. Během expozice jsme každých 24 hodin pomocí přenosné fluorescenční kamery Handy Fluor Cam 010 (PSI, ČR) měřili parametry indukované fluorescence chlorofylu (FV/FM, II a NPQ). Výsledky měření ukázaly, že experimentální přídavek 32mM ribitolu vedl ke zvýšení hodnot FV/FM. Toto zvýšení však bylo patrné pouze při teplotě –5 °C, u ostatních experimentálních teplot (0 °C a +5 °C) nebyly zjištěny žádné výrazné rozdíly ve srovnání s kontrolou (tj. stélky v destilované vodě, bez přídavku ribitolu). To ukazuje, že pozitivní efekt externě dodaného ribitolu na primární procesy fotosyntézy v lišejnících se objevuje pouze za mrazových teplot. Tento účinek tak může souviset s kryoprotektivní rolí polyolů v rostlinách a symbiotických organismech (lišejnících). Přídavek 32mM ribitolu se také pozitivně projevil ve zvýšení hodnot kvantového výtěžku fotochemických procesů v PS II ( II) za všech experimentálních teplot. 5 Naproti tomu přídavek 50mM ribitolu měl v porovnání s kontrolou negativní účinek na FV/FM, II za všech sledovaných experimentálních teplot. Výraznější vlivy ve variantě AR nebyly pozorovány. Klíčová slova: lišejníky, polyoly, ribitol, manitol, arabitol, nízké a mrazové teploty, Xanthoparmelia somloensis (Gyelnik) Hale, Trebouxia 6 Abstract My bachelor thesis was composed of two basic parts. The first one was focused on a literature review on polyols (sugar alcohols), their role in lichen protection against low and freeze temperatures. The second part of my bachelor thesis was an experimental work. The main aim of the diploma thesis was to find effective concentrations of polyols, ribitol in particular, in mechanisms of lichen resistance to low and freezing temperatures. Thalli of foliose lichen species Xanthoparmelia somloensis (Gyelnik) Hale, with Trebouxia as a photobiont, were collected close to the Brno Lake dam, Czech Republic, in November 2006. The thalli were cleaned with deionized water. Then, a half of the selected segments were subjected to acetone rinsing. In acetone rinsed (AR) lichen segments, majority of secondary metabolites were extracted. Untreated (non-AR) and acetone-rinsed (AR) thalli were exposed to 32mM (5 g/l) or 50mM (7.8 g/l) ribitol for 10 min, left in closed Petri dishes under dim light for 24 h, and then exposed to the respective ribitol concentration for 10 min again. Thalli segments of control (C), non- AR and AR treatments were then closed into Petri dishes using Parafilm and cultivated at -5, 0, and +5 °C (LABIO, Prague, Czech Republic) under 300 μmol.m-2.s-1 of photosynthetically active radiation for 168 h. Every 24 h, parameters of chlorophyll fluorescence (potential yield of photochemical reactions in PS II – FV/FM, effective quantum yield of photochemical reactions in PS II – II and non-photochemical quenching of chlorophyll fluorescence – NPQ) were measured using a HFC-010 portable fluorometer (Photon Systems Instruments, Brno, Czech Republic). During cultivation, the Petri dishes were opened very shortly and lichen thalli were sprayed several times by demineralised water. In all the examined thalli segments, an apparent drop in potential yield of photochemical reactions in PS II (FV/FM) values was found after 24-h exposure to respective temperature. Maximum decrease in FV/FM was found at -5 °C, while the decrease was much less pronounced at 0 and +5 °C. Lower FV/FM values were usually found in control thalli than in acetone rinsed thalli segments (AR) and in non-AR. Application of ribitol affected FV/FM values only at -5 °C. While positive effect on FV/FM was apparent in thalli treated with 32mM ribitol, negative effect occurred when treated with 50mM ribitol. 7 Effective quantum yield of photochemical reactions in PS II ( II) showed similar response to temperature treatments as FV/FM. Absolute II values decreased with treatment temperature. Positive effect of 32 mM ribitol on II was found at -5 °C in AR thalli while no such response was apparent in non-AR thalli. Key-words: lichens, polyols, sugar alcohols, ribitol, mannitol, arabitol, low and freezing temperatures, Xanthoparmelia somloensis (Gyelnik) Hale, Trebouxia 8 OBSAH: 1. ÚVOD 1 2. PŘEHLED LITERATURY 5 2.1. Lišejníky 5 2.1.1. Anatomická a morfologická stavba lišejníků 5 2.2. Mechanismy kryoprotekce 9 2.2.1. Vliv nízkých teplot na rozšíření rostlin na Zemi 9 2.2.2. Teplotní stres 10 2.2.2.1. Chladový stres 10 2.2.2.2. Mrazový stres 11 2.2.3. Proces mrznutí 11 2.2.3.1. Nukleace ledu 12 2.2.3.2. Antifreeze proteins AFP´s 12 2.2.4. Mechanismy mrazové odolnosti 14 2.2.4.1. Mrazová avoidance 14 2.2.4.2. Mrazová tolerance 14 2.2.5. Signální dráhy v buňkách ovlivněných nízkou teplotou 15 2.2.5.1. Reakce vyvolané pomalým poklesem teploty 15 2.2.5.2. Reakce vyvolané rychlým poklesem teploty 16 2.3. Kryoprotekce a kryoprotektanty 19 2.4. Polyoly 21 2.4.1. Obecná charakteristika polyolů 21 2.4.2. Biologické vlastnosti polyolů 24 2.4.2.1. Výskyt, funkce a použití 24 2.4.3. Přirozená množství polyolů v lišejnících 26 2.4.4. Metabolismus polyolů 28 3. MATERIÁL A METODY 31 3.1. Charakteristika experimentálních druhů 31 3.2. Cíle experimentu 36 3.3. Průběh přípravy experimentu a expozice stélek 37 3.4. Měření parametrů indukované fluorescence chlorofylu 39 3.5. Stanovení obsahu chlorofylů a karotenoidů 40 3.6. Stanovení obsahu polyolů 41 9 4. VÝSLEDKY 42 4.1. Xanthoparmelia somloensis 42 4.1.1. FV/FM 42 4.1.2. 42 4.1.3. NPQ 43 4.2. Cetraria islandica 43 4.2.1. FV/FM 43 4.2.2. 43 4.2.3. NPQ 44 4.3. Flavocetraria nivalis 44 4.3.1. FV/FM 44 4.3.2. 44 4.3.3. NPQ 45 4.4. Obsah chlorofylů a karotenoidů 45 4.5. Obsah polyolů 45 5. DISKUZE 50 6. ZÁVĚR 53 7. SEZNAM CITOVANÉ LITERATURY 55 8. PŘÍLOHY 65 10 Přehled používaných symbolů a zkratek: DW - suchá hmotnost (Syn: hmotnost sušiny) stélky F0 - minimální výtěžek fluorescence chlorofylu změřený na zatemněných vzorcích
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