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Reprodukční Chování a Pohlavní Výběr U Hořavek (Acheilognathinae) S Důrazem Na Hořavku Duhovou (Rhodeus Amarus)

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Reprodukční Chování a Pohlavní Výběr U Hořavek (Acheilognathinae) S Důrazem Na Hořavku Duhovou (Rhodeus Amarus) MASARYKOVA UNIVERZITA V BRNĚ PŘÍRODOVĚDECKÁ FAKULTA ÚSTAV BOTANIKY A ZOOLOGIE Reprodukční chování a pohlavní výběr u hořavek (Acheilognathinae) s důrazem na hořavku duhovou (Rhodeus amarus) Bakalářská práce Brno 2008 Autor: Radomil Řežucha MASARYKOVA UNIVERZITA V BRNĚ PŘÍRODOVĚDECKÁ FAKULTA ÚSTAV BOTANIKY A ZOOLOGIE Reprodukční chování a pohlavní výběr u hořavek (Acheilognathinae) s důrazem na hořavku duhovou (Rhodeus amarus) Bakalářská práce Autor: Radomil Řežucha Brno 2008 Vedoucí BP: RNDr. Martin Reichard, PhD. Prohlášení Souhlasím s uložením této bakalářské práce v knihovně Ústavu botaniky a zoologie PřF MU v Brně, případně v jiné knihovně MU, s jejím veřejným půjčováním a využitím pro vě- decké, vzdělávací nebo jiné veřejně prospěšné účely, a to za předpokladu, že převzaté infor- mace budou řádně citovány a nebudou využívány komerčně. Datum …………………….. ……………………………. Radomil Řežucha Poděkování Chtěl bych poděkovat RNDr. Martinu Reichardovi, Phd. za pomoc a podporu při zpraco- vání této práce. Abstrakt Cílem této práce je shrnout poznatky o reprodukčním chování hořavek (Acheilognathi- nae), hlavně pak hořavky duhové (Rhodeus amarus), a charakterizovat mechanizmy pohlav- ního výběru, na něj navazující párovací systémy a alternativní párovací taktiky. Pohlavní výběr, tedy ústřední téma této práce, zvýhodňuje určitého jedince před ostatní- mi jedinci stejného pohlaví a dělí se na intrasexuální a intersexuální. Intersexuální pohlavní výběr podléhá samičím preferencím, které se dají vysvětlit Fisherovou hypotézou, hypotézou dobrých genů či využitím smyslových vjemů. Z evolučního hlediska je pro samičky nezbytné maximalizovat svůj přímý a nepřímý zisk. Přímý zisk ovlivňuje samičku samotnou a může být ve formě rodičovské péče, svatebních darů, většího počtu oplodněných vajíček či kvalit- ního místa pro rozmnožování. Nepřímý zisk ovlivňuje potomstvo samičky prostřednictvím kvalitních genů partnera a samička se pro jeho maximalizaci může řídit množstvím signálů samečkovy kvality, např. akustickými, vizuálními a chemickými signály. Párovací systémy jsou ovlivněny možnostmi získání partnera buď přímo, nebo pomocí vlastnictví zdrojů nutných pro úspěšné rozmnožování. Dělí se na monogamii, polygamii (ta dále na polygynii a polyandrii) a promiskuitu (polygynandrii). Na párovací systémy se váže i vznik alternativních rozmnožovacích taktik, které mohou být podmíněny populační hustotou, velikostí, sociálním statusem nebo kondicí samečka či přímo fixováno v jeho genotypu. Hořavky (Acheilognathinae) jsou skupinou ryb s neobvyklým způsobem rozmnožování, kdy kladou jikry do žaberní dutiny mlžů čeledí Unionidae a Margaritiferidae. Slouží jako mo- delový organismus nejen pro výzkum reprodukčního chování, ale i pro výzkum koevoluce. Jsou zastoupeny hlavně v Asii, ale i v Evropě a to jediným druhem, hořavkou duhovou (Rho- deus amarus). Samečci se vyznačují využíváním alternativních taktik při rozmnožování, které vychází z jejich velikosti, úrovně dominance a populační hustoty. Samičí volba je v počátku ovlivněna barvou samce a intenzitou námluv a finální rozhodnutí vychází z kvality mlže. V současné době je celosvětovým problémem sílící eutrofizace a s ní spojený nárůst sinic a řas. Bude jistě přínosné doplnit současný sporý počet studií o vlivu snížené viditelnosti ve vodním prostředí na pohlavní výběr o experimenty s hořavkou duhovou (Rhodeus amarus). Abstract Reproductive behaviour and sexual selection in bitterling fishes (Acheilo- gnathinae) with a special emphasis on the European bitterling (Rhodeus amarus). Aim of this thesis is to summarize findings about reproductive behaviour of bitterlings (Acheilognathinae), especially the European bitterling (Rhodeus amarus), and to characterize mechanisms of sexual selection, types of mating systems and nature of alternative mating tactics. Sexual selection, a central subject of this thesis, gives advantage to one individual over another of the same sex and can be divided into intrasexual and intersexual. Intersexual se- lection is subject to female preference, that can be explained by Fisher‘s hypothesis, hypothe- sis of good genes and by sensory exploitation. From evolutionary perspective it is necessary for females to maximalize their direct and indirect benefit. Direct benefit can be in a form of parental care, nuptial gifts, bigger amount of fertilized eggs or a quality site for mating and it has impact on female herself. Indirect benefit affects offspring of female through high-quality male genes and female can employ number of signals of male quality to maximize it, e. g. acoustic, visual and chemical signals. Mating systems are based on ability to monopolize mate directly, or through possession of resources essential for successful breeding. It can be divided into monogamy, polygamy (then to polygyny and polyandry) and promiscuity (polygynandry). Origin and evolution of alternative mating tactics is related to mating systems. They can be influenced by density, social status, male condition and size or fixed in genotype. Bitterlings (Acheilognathinae) are a group of freshwater fish with unusual way of repro- duction as they spawn on the gills of Unionid and Margaritiferid mussels. They serve as a model species not only for a research of reproductive behaviour, but also for research of co- evolution. They are present mainly in Asia and one species, Rhodeus amarus, occurs in Euro- pe. Males are characterized by using of alternative mating tactics, that are affected by male size, dominance and density. Female choice is initially influenced by male‘s colour and court- ship vigour while final decision depends on mussel quality. Increasing eutrophication and accompanying grow of cyanobacteria and algae is problem worldwide. It would be beneficial to supply current insufficient amount of studies about in- fluence of water turbidity on sexual selection by experiments with the European bitterling (Rhodeus amarus). Obsah 1 ÚVOD .................................................................................................................................................. 1 2 POHLAVNÍ VÝBĚR ...................................................................................................................... 2 2.1 ANISOGAMIE ................................................................................................................................... 2 2.2 DRUHY A MECHANIZMY POHLAVNÍHO VÝBĚRU ........................................................................ 2 2.2.1 Intrasexuální výběr ............................................................................................. 3 2.2.2 Intersexuální výběr ............................................................................................. 3 2.2.2.1 Samičí preference ................................................................................................. 3 2.2.2.2 Samičí volba (přímý × nepřímý zisk) ..................................................................... 5 3 PÁROVACÍ SYSTÉMY ...............................................................................................................10 3.1 ALTERNATIVNÍ TAKTIKY..............................................................................................................13 4 HOŘAVKY (ACHEILOGNATHINAE) .................................................................................16 4.1 HOŘAVKA DUHOVÁ (RHODEUS AMARUS)...................................................................................17 4.1.1 Charakteristika .................................................................................................. 17 4.1.2 Reprodukční chování ........................................................................................ 18 4.1.2.1 Pohlavní výběr ............................................................................................ 19 4.1.3 Metodika výzkumu ........................................................................................... 22 5 DISKUZE ..........................................................................................................................................24 6 POUŽITÉ ZDROJE ......................................................................................................................26 7 OBRÁZKY .......................................................................................................................................31 ÚVOD 1 1 Úvod Již Charles Darwin si všiml, že evoluci některých struktur, např. rohů, paroží, extrava- gantních per či ploutví, nápadného zbarvení a opeření, nebo jiných ornamentů (tzv. sekundár- ních pohlavních znaků), které přímo neslouží k přežití jedince, lze jen stěží vysvětlit mecha- nismy přírodního výběru. Zavedl proto koncept výběru pohlavního, ve kterém rozeznával dvě složky: intrasexuální samčí souboj o samice a intersexuální samičí volbu samce. Položil tak základy současného myšlení a přístupu ke studiu pohlavního rozmnožování organismů (AN- DERSSON 1994). Na jeho práci se snaží navázat současný výzkum, který se zabývá intenzi- tou a mechanismy pohlavního výběru u modelových organismů, které jsou k tomuto studiu vhodné, ať už z behaviorálního či fyziologického hlediska. Vhodné kandidáty nalézáme hlav- ně v řadách hmyzu a ptáků, jakožto nejlépe prostudované skupiny obratlovců (AMUNDSEN 2006), v menší míře pak u dalších skupin živočichů. Jedním z řady nabízejících se taxonů je díky svému epigamnímu (reprodukčnímu) cho- vání i skupina hořavek, drobných ryb z čeledi Cyprinidae (podčeleď Acheilognathinae), která se vyskytuje hlavně ve východní Asii a u nás
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