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MASARYKOVA UNIVERZITA Analýza Vybraných Ekologických a Evolučně-Behaviorálních Aspektů V Systému Parazit-Hostitel

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MASARYKOVA UNIVERZITA Analýza Vybraných Ekologických a Evolučně-Behaviorálních Aspektů V Systému Parazit-Hostitel MASARYKOVA UNIVERZITA Přírodovědecká fakulta Martina DÁVIDOVÁ Analýza vybraných ekologických a evolučně-behaviorálních aspektů v systému parazit-hostitel Disertační práce Školitel: doc. RNDr. Milan Gelnar, CSc. Brno, 2008 Bibliografická identifikace Jméno a příjmení autora: Martina Dávidová Název disertační práce: Analýza vybraných ekologických a evolučně-behaviorálních aspektů v systému parazit-hostitel Název disertační práce anglicky: Analysis of selected ecological and evolutionary-behavioural aspects of host-parasite systems Studijní program: Biologie Studijní obor (směr), kombinace oborů: Parazitologie Školitel: doc. RNDr. Milan Gelnar, CSc. Rok obhajoby: 2008 Klíčová slova v češtině: systém parazit-rybí hostitel, interakce, biotické a abiotické faktory, virulence, selektivní predace, pohlavní výběr Klíčová slova v angličtině: host fish-parasite system, interactions, biotic and abiotic factors, virulence, selective predation, sexual selection © Martina Dávidová, Masarykova univerzita, Brno, 2008 Poděkování Mé poděkování patří mému školiteli doc. RNDr. Milanu Gelnarovi, CSc. za poskytnutí zázemí, rad, trpělivosti a finanční podpory při zpracování této práce. Můj velký dík určitě patří Andree Šimkové, Markétě Ondračkové a Radimu Blažkovi za jejich podporu, pomoc a optimismus v každé situaci; Jirkovi Jarkovskému, Honzovi Mužíkovi a Katce Houdkové za pomoc při řešení statistických zapeklitostí; Ivetě Matějusové za korektury anglicky psaného textu; Pavlu Jurajdovi, Martinu Reichardovi, Michalu Janáčovi a Zdeničce Valové za jejich neocenitelnou pomoc v terénu a také všem z oddělení parazitologie, kteří se jakýmkoliv způsobem podíleli na sběru dat. Prof. Arnemu Skorpingovi děkuji za možnost vyzkoušet si „něco jiného“ na univerzitě v norském Bergenu. Především děkuji své rodině a přátelům za podporu a pochopení, které jsem často potřebovala. Abstrakt Studium vybraných interakcí v systému parazit-hostitel (model: ryba-mnohobuněčný cizopasník) bylo ústředním tématem předložené disertační práce. Práce je tvořena sedmi publikacemi a podle úhlu pohledu na studium komplexu parazit-hostitel rozdělena na dvě části, které hodnotí dva různé aspekty uvedeného systému: (1) ekologický a (2) evolučně- behaviorální. V první části práce byl vyhodnocen vliv vybraných biotických a abiotických faktorů na strukturu a složení společenstev parazitů. Modelovým hostitelem těchto tří studií byla drobná sladkovodní ryba hořavka duhová (Rhodeus amarus), charakteristická širokým areálem rozšíření a unikátním typem rozmnožování. Sezonní změny teploty vody kromě vlivu na dynamiku výskytu a epidemiologii parazitů významně ovlivňovaly i metrickou variabilitu sklerotizovaných struktur jejich příchytného aparátu (článek A), také charakter prostředí hostitele signifikantně ovlivnil složení společenstev cizopasníků (článek B), dále byl potvrzen úzký vztah mezi potravní specializací hostitele a zastoupením především endoparazitických druhů (článek B a C). Nakonec byl potvrzen i vztah mezi velikostí (věkem) hostitele a složením společenstva parazitů (článek B), ačkoliv variabilita vysvětlená délkou byla velmi nízká. Druhá, převážně na experimentálních studiích založená část práce byla zaměřena na studium vlivu parazitů na imunokompetenci, chování a reprodukci hostitele. V této části byly použity čtyři parazito-hostitelské systémy, přičemž každý systém byl specifický pro daný typ studie. Hypotéza pozitivní závislosti plodnosti a virulence (míra poškození hostitele) parazita byla testovaná v experimentálních podmínkách na modelu lososa obecného (Salmo salar) a parazitického korýše Lepeophtheirus salmonis (článek D). Získané výsledky naznačují, že hierarchické postavení subdominantních ryb, které je úzce spojené se sociálním stresem, může signifikantně ovlivnit výsledky měření virulence parazita. Také výsledky experimentálních testů zaměřených na sledování selektivní predace karase stříbřitého (Carassius auratus), infikovaného larválními stádii strigeidní motolice Posthodiplostomum cuticola, okounem říčním (Perca fluviatilis) podporují hypotézu o potenciálním vlivu parazita na chování hostitele (článek E). Role parazitů při pohlavním výběru byla v našem případě testovaná na dvou odlišných parazito-hostitelských modelech. Pozitivní vztah mezi sexuální ornamentací a parazitární infekcí byl potvrzen u cejna velkého (Abramis brama), kdy samci v období tření investovali více energie do reprodukce (rozvoj sexuální ornamentace) než do obranyschopnosti proti parazitům (článek G), avšak žádný vztah mezi sexuální ornamentací (zbarvením oka) a parazitární infekcí nebyl zjištěn u samců hořavky duhové (R. amarus) (článek F). Abstract The thesis summarizes analyses of the selected processes and patterns of host-parasite interactions in the model of fish-parasite systems. The thesis is composed of 7 papers divided in the two sections: (1) ecological aspects of the host-parasite system and (2) evolutionary- behavioural aspects of the host-parasite systems. The European bitterling (R. amarus), small cyprinid fish inhabiting lentic and riverine waters with an unusual spawning relationship with freshwater mussels, is a model species for ecological studies and was used in studies included in the first section. Four specific host-parasite systems were used in the second part of the work, which was mainly experimental. The first section of the thesis was aimed to analyze the role of selected abiotic and biotic factors in the parasite infections. It was found that the occurrence of parasites and metrical variability of their attachment apparatus depended on seasonal changes in water temperature (attachment A); the composition and structure of parasite communities was significantly affected by the host habitat (attachment B); the effect of host-specific diet on the structure primary endoparasite communities was confirmed (attachments B and C) and finally host body size (age) and the structure of parasite communities were weakly associated (attachment B). The second part of the thesis was principally aimed to investigate the effect of metazoan parasites on the immunocompetence, behaviour and reproduction of the host fish. The hypothesis of positive correlation between fecundity and virulence (disease severity) of parasites was tested using the ectoparasitic crustacean Lepeophtheirus salmonis and the host Salmo salar in the experimental conditions (attachment D). The results of this experiment indicate that fish status linked with a social stress of subordinate fish can significantly affect the virulence measurements. Experimental tests on the selective predation rate of the Posthodiplostomum cuticola infected fish (Carassius auratus) by non-host predator (Perca fluviatilis) supported the hypothesis of the effect of parasite on the fish behaviour (attachment E). The relationship between expression of the sexual ornamentation and parasite infection was studied on two independent models of freshwater fishes (Abramis brama and R. amarus). Males of A. brama that allocated more energy resources into reproduction by developing more intensive sexual ornamentation were more susceptible to the metazoan ectoparasite infection (attachment G). On the other hand the relationship between sexual ornamentation and parasite infection rate was not confirmed in the case of males of R. amarus (attachment F). The results offered several insights into complex host-parasite relationships and provided several tests of evolutionary hypotheses. OBSAH 1 Úvod 8 2 Cíle práce a přehled publikací 9 3 Literární přehled 10 3.1 Úrovně studia parazitů 11 3.2 Distribuce cizopasníků 13 3.3 Diverzita a druhová bohatost cizopasníků 15 3.3.1 Faktory ovlivňující strukturu společenstev cizopasníků ryb 15 3.3.1.1 Biotické faktory 16 3.3.1.2 Abiotické faktory 18 3.3.1.3 Ostatní faktory 19 3.4 Strategie využívání hostitele parazitem 20 3.4.1 Složky životních historií parazita („Life-history traits“) 20 3.4.1.1 Velikost parazita 21 3.4.1.2 Věk a rychlost vývoje parazita 21 3.4.1.3 Produkce potomků 21 3.4.2 Evoluce virulence 22 3.4.3 Vliv parazita na chování hostitele 24 3.4.4 Role parazitů při pohlavním výběru hostitele 27 4 Materiál a metody 29 4.1 Lokality výzkumu a studované systémy 29 4.2 Fixace a determinace parazitologického materiálu 29 4.3 Epidemiologické charakteristiky a klasifikace parazitů 30 4.4 Zpracování dat 30 4.4.1 Biotické a abiotické faktory ovlivňující parazitární infekci 30 4.4.2 Analýza vlivu parazita na rybího hostitele 31 5 Výsledky a závěry 39 5.1 Role vybraných biotických a abiotických faktorů na strukturu společenstev 39 parazitů hořavky duhové (Rhodeus amarus) 5.2 Stanovení vlivu parazita na rybího hostitele 41 6 Shrnutí a další perspektivy výzkumu 44 7 Použitá literatura 46 8 Publikace tvořící disertační práci 59 1 Úvod 1 Úvod Paraziti tvoří převážnou většinu na Zemi žijících organismů (Poulin a Morand 2000). Jejich funkce jako speciálních konzumentů a také nesporný vliv na biodiverzitu je staví do role významných prvků v mnoha ekosystémech. Lze říci, že teorie dualismu je v ekologii parazitismu dominantní: paraziti mohou generovat diverzitu volně žijících organismů a současně vést k jejich extinkci, mohou kastrovat svého hostitele a také způsobovat jeho růst, mohou stimulovat imunitní odpověď a zároveň podporovat rozvoj sekundární chronické infekce (Hudson 2005). Paraziti vstupují do interakce se svým hostitelem na několika úrovních, od boje mezi imunitní odpovědí hostitele a parazitem na molekulární úrovni až po strukturu volně žijících společenstev. Parazitické organismy mají vliv nejenom na populační
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