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Bc Prace Cely Text 23 04 13 Finally3 MASARYKOVA UNIVERZITA PŘÍRODOVĚDECKÁ FAKULTA ÚSTAV BOTANIKY A ZOOLOGIE Bakalářská práce Brno 2013 Jana Přibylová MASARYKOVA UNIVERZITA PŘÍRODOVĚDECKÁ FAKULTA ÚSTAV BOTANIKY A ZOOLOGIE Bakteriální endocytobionti voln ě žijících améb: synopse druh ů determinovaných s podporou molekulárních marker ů Bakalářská práce Jana Přibylová Vedoucí práce: prof. MVDr. Iva Dyková, DrSc. Brno 2013 Bibliografický záznam Autor: Bc. Jana Přibylová Přírodovědecká fakulta, Masarykova univerzita Ústav botaniky a zoologie Bakteriální endocytobionti volně žijících améb: Název práce: synopse druhů determinovaných s podporou molekulárních markerů Studijní program: Ekologická a evoluční biologie Studijní obor: Ekologická a evoluční biologie Vedoucí práce: prof. MVDr. Iva Dyková, DrSc. Akademický rok: 2012/2013 Počet stran: 34 Klíčová slova: Volně žijící améby, amfizoické améby, bakteriální symbionti, endocytobionti, mikroorganismy odolávající natrávení améb Bibliographic Entry Author Bc. Jana Přibylová Faculty of Science, Masaryk University Department of Botany and Zoology Bacterial endocytobionts of free-living amoebae: Title of Thesis: synopsis of species identified with the aid of molecular markers Degree programme: Ecological and evolutionary biology Field of Study: Ecological and evolutionary biology Supervisor: prof. MVDr. Iva Dyková, DrSc. Academic Year: 2012/2013 Number of Pages: 34 Keywords: Free-living amobeae, amphizoic amoebae, bacterial symbionts, endocytobionts, amoeba-resisting microorganisms Abstrakt Bakalá řská práce je zam ěř ena na nejd ůležit ější poznatky o symbiotickém vztahu voln ě žijících améb a prokaryotických organism ů. Pomocí molekulárních metod byli v amébách do sou časnosti identifikováni symbiondocytobionti hlavn ě ze skupin proteobakterií, rickettsií a chlamydií. Práce obsahuje seznam popsaných symbiont ů/endocytobiont ů, který je dopln ěn krátkým souhrnem dosud získaných informací o jejich intracelulárním zp ůsobu života. Abstract Presented is an overview of the most important data published recently on symbiotic associations of free-living amoebae and prokaryotic organisms. The main emphasis is on proteobacteria, rickettsiae and chlamydiae that have been determined as symbionts/endocytobionts of free-living amoebae, using molecular methods. Synopsis of identified symbionts/endocytobionts of free-living amoebae and relevant references are supplemented with a short analysis of the current state of knowledge in the field. Kopie zadání Poděkování Děkuji vedoucí bakalářské práce prof. MVDr. Ivě Dykové, DrSc. za odborné vedení, ochotu a trpělivost při zpracovávání textu. Poděkování patří také rodině a přátelům za podporu. Prohlášení Prohlašuji, že jsem svoji bakalářskou práci vypracovala samostatně s využitím informačních zdrojů, které jsou v práci citovány. Brno 2. května 2013 ……………………………… Jana Přibylová Obsah 1 Úvod ............................................................................................................................. 10 1.1 Cíle práce ......................................................................................................... 10 1.2 Vysv ětlení pojm ů ............................................................................................. 10 1.2.1 Voln ě žijící améby .................................................................................... 10 1.2.2 Symbióza a endocytobióza obecn ě ........................................................... 11 1.2.3 Endocytobióza .......................................................................................... 12 1.2.4 Voln ě žijící améby jako rezervoáry patogen ů .......................................... 13 1.2.5 Endosymbionti nepatogenních voln ě žijících améb ................................. 14 1.2.6 Fyziologické interakce voln ě žijících améb a endocytobiont ů ................. 15 1.2.7 Genetický pohled na symbiózu/endocytobiózu ........................................ 16 2 Symbióza/endocytobióza voln ě žijících améb a prokaryot ů ...................................... 17 2.1 Symbióza/endocytobióza voln ě žijících améb a proteobakterií ....................... 17 2.2 Symbióza/endocytobióza voln ě žijících améb a rickettsií ............................... 19 2.3 Symbióza/endocytobióza voln ě žijících améb a chlamydií ............................. 20 3 Seznam symbiont ů/ endocytobiont ů voln ě žijících améb ........................................... 22 4 Souhrn ......................................................................................................................... 26 5 Seznam použité literatury ........................................................................................... 28 1 Úvod 1.1 Cíle práce Cílem rešeršní bakalá řské práce bylo získání p řehledu o sou časném stavu poznání symbiotických asociací dvou skupin mikroorganism ů, voln ě žijících/amfizoických améb a bakterií. Zadání práce bylo zám ěrn ě omezeno na rešerši literárních údaj ů podložených molekulárními daty. Téma navázalo na výsledky výzkumu voln ě žijících améb v roli potenciálních patogen ů ryb, jehož sou částí bylo i studium ultrastruktury izolovaných kmen ů. Transmisní elektronová mikroskopie prokázala p řítomnost bakterií v cytoplazm ě amébových trofozoitů. Krom ě obvyklé lokalizace bakterií ve fagosomech, byla u velkého po čtu kmen ů zaznamenána i lokalizace v symbiosomech (Dyková & Kostka 2013). Z této skute čnosti vyplynula snaha p řipravit se na podrobn ější studium tematiky symbiotických vztah ů améb a bakterií v četn ě jejich determinace. 1.2 Vysv ětlení pojm ů 1.2.1 Voln ě žijící améby Améby pat ří mezi eukaryotické organismy. Podle obecného konsensu termín "amoeba" vyjad řuje typ bun ěč né organizace a není ozna čením pro taxon (Smirnov & Brown 2004). Existují dv ě významné skupiny améb, které se liší energetickým metabolismem. Jsou to endozoické a parazitické mikroaerofilní améby st řevního traktu, k nimž pat ří mj. i p ůvodce amébové dyzentérie a jaterních absces ů u člov ěka ( Entamoeba histolytica ), a voln ě žijící, aerobní améby. Voln ě žijící améby jsou často ozna čovány za kosmopolitn ě rozší řené eukaryotické organismy. Žijí ve v ětšin ě vodních a suchozemských biotop ů, v půdě, sedimentech, na částicích suspendovaných ve vod ě, a také jako epi- a endobionti bezobratlých a obratlovc ů. Voln ě žijící améby nejsou primárn ě uzp ůsobeny k parazitismu. Nesnaží se udržet svého hostitele naživu, jelikož na n ěm nejsou reproduk čně závislé. Jsou schopny p řežívat uvnit ř hostitele, v n ěmž mohou zahájit parazitický zp ůsob života a stát se etiologickým agens (Schuster & Visvesvara 2004). Pro voln ě žijící améby, které se mohou množit ve vn ějším prost ředí i v hostitelích je 10 používán termín amfizoické améby (Page 1974). K amfizoickým amébám pat ří i druhy, které jsou významnými patogeny pro člov ěka, tj. p ůvodci primární amébové meningoencephalitidy (PAM), granulomatózní amébové encephalitidy (GAE) a p ůvodci amébových keratitid (AK). Diverzita voln ě žijících améb je dokumentována velkým po čtem popisných prací publikovaných zejména v minulém století. Schopnost amfizoické (exo- a endozoické) existence t ěchto organism ů a patogenita prokázaná u n ěkterých druh ů voln ě žijících améb obrací pozornost biolog ů k voln ě žijícím amébám více než 50 let. V tradi ční, na morfologii založené klasifikaci, byly voln ě žijící améby řazeny do spole čné skupiny Sarcodina (Pawlowski 2008) více než 100 let. K poznání voln ě žijících améb zásadním zp ůsobem p řisp ěl Frederik C. Page, který do v ětšiny popis ů jednotlivých taxon ů za řadil i detaily jejich ultrastruktury (Page 1980, 1983, 1988) a vytvo řil základ pro moderní hodnocení jednotlivých morfotyp ů améb (Page 1991). Další vývoj poznání, pozitivn ě ovlivn ěný vstupem molekulárních a fylogenetických metod, přinesl výrazné zm ěny v názorech na systematiku voln ě žijících améb. Dnes víme, že tyto organismy pat ří do 5 z 6 navržených superskupin eukaryotických organism ů (Adl et al. 2005; Pawlowski 2008). Velké úsilí je v sou časné dob ě v ěnováno poznávání fylogenetických vztah ů na úrovni rod ů. 1.2.2 Symbióza a endocytobióza obecn ě Ačkoliv slovo symbióza často navozuje p ředstavu vzájemn ě výhodného soužití dvou organism ů, v obecné rovin ě je definována jako soužití nebo asociace rozdílných biont ů. Pokud je termín symbióza použit v širokém smyslu slova, zahrnuje r ůzné úrovn ě asociací jednotlivých organism ů definované jako foréza, komensalismus, mutualismus a parazitismus (Smith & Douglas 1987). Definice symbiózy v užším smyslu slova zd ůraz ňuje dlouhotrvající nebo permanentní soužití organism ů a zahrnuje blíže nedefinované, mutualistické a parazitické asociace. Asociované organismy mohou v rámci spole čné koexistence získat prosp ěch, být poškozovány, nebo se nemusí ovliv ňovat. Charakter vzájemného ovlivn ění symbiont ů nemusí být vzhledem ke zm ěnám ekologických podmínek či ontogenetických stadií organismů ur čen vždy jednozna čně. 11 Větší z asociovaných organism ů je často ozna čován za hostitele menšího symbionta. Symbiont je s hostitelem vždy ve fyzickém kontaktu. V případ ě obligátní symbiózy není symbiont schopen bez hostitele p řežít nebo se množit a k tomuto zp ůsobu života se obvykle p řizp ůsobuje modifikací své morfologie i biologie. Symbiont si svého hostitele vybírá v procesu rozpoznávání. Množství hostitel ů, se kterými je symbiont schopen soužití, záleží na stupni hostitelské specifity. Krom ě specifity hostitelské, je specifita symbiózy rozd ělována na ekologickou a druhovou. Druhová specifita neuvažuje ekologické podmínky. Spojení symbionta
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