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VLIV TERMOREGULAČNÍHO CHOVÁNÍ SAMIC ČOLKŮ NA FENOTYP POTOMSTVA Diplomová Práce

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VLIV TERMOREGULAČNÍHO CHOVÁNÍ SAMIC ČOLKŮ NA FENOTYP POTOMSTVA Diplomová Práce MASARYKOVA UNIVERZITA PŘÍRODOVĚDECKÁ FAKULTA ÚSTAV BOTANIKY A ZOOLOGIE VLIV TERMOREGULAČNÍHO CHOVÁNÍ SAMIC ČOLKŮ NA FENOTYP POTOMSTVA Diplomová práce Eliška Toufarová VEDOUCÍ PRÁCE: Mgr. Lumír Gvoždík, Ph.D. BRNO 2013 Bibliografický záznam Autor: Bc. Eliška Toufarová Přirodovědecka fakulta, Masarykova univerzita Ústav botaniky a zoologie Vliv termoregulačního chování samic čolků na fenotyp Název práce: potomstva Studijní program: Biologie Studijní obor: Ekologická a evoluční biologie Vedoucí práce: Mgr. Lumír Gvoždík, Ph.D. Akademický rok: 2013 Počet stran: 50 Klíčová slova: Reprodukce, termoregulace, mateřský efekt, čolek horský, fenotyp, potomstvo Bibliographic Entry Author Bc. Eliška Toufarová Faculty of Science, Masaryk University Department of botany and zoology Influence of thermoregulatory in female newts on Title of Thesis: offspring phenotype Degree programme: Biology Field of Study: Field of Study Supervisor: Mgr. Lumír Gvoždík, Ph.D. Academic Year: 2013 Number of Pages: 50 Keywords: Reproduction, thermoregulation, maternal effect, alpine newt, phenotype, offspring ABSTRAKT Reproduktivní samice mnoha ektotermů si udržují teploty těla, které se liší od jejich teplot těla mimo období rozmnožování. Tato změna bývá vysvětlována ve shodě s hypotézou mateřské manipulace, tj. matka mění chování, aby zvýšila fitness svého potomstva. Cíle této práce byly dva. (1) Zjistit, zda reprodukce ovlivňuje termoregulační chování samic u čolka horského, Ichthyosaura alpestris; (2) zjistit vliv rozdílných teplotních nároků reproduktivních a nereproduktivních samic na fenotyp potomstva. Výsledky ukázaly, že kladoucí samice si udržovaly nižší a méně proměnlivé teploty těla než nereproduktivní. To potvrzuje dřívější tvrzení, že ke změně termoregulačního chování dochází dokonce i u druhů, u kterých samice kladou vejce v raných stádiích embryogeneze. Teplotní režimy simulující teploty těla reproduktivních a nereproduktivních samic měly neprůkazný vliv na velikost vajec, rychlost vývoje a velikost při vylíhnutí. Tyto výsledky naznačují, že termoregulační chování samic se vyvinulo dříve než pokročilé způsoby rozmnožování a tento fenomén u čolků není vysvětlitelný mateřskou manipulací fenotypů potomstva. Zdá se pravděpodobné, že reproduktivní samice modifikují termoregulační chování, aby maximalizovaly jejich rychlost kladení vajec. ABSTRACT Reproductive females of many ectotherms maintain body temperatures that differ from their body temperatures during non-reproductive period. This change has been explained in accordance with the maternal manipulation hypothesis, i.e., mother changes behavior to increase her offspring fitness. The aim of this study was twofold. (1) To determine whether reproduction affects female thermoregulatory behavior in the Alpine newt, Ichthyosaura alpestris; (2) to determine the influence of different thermal requirements of reproductive and non-reproductive females on offspring phenotype. The results showed that ovipositing females maintained lower and less variable body temperatures than non-reproductive ones. It corroborates the previous claim that the change in thermoregulatory behavior occurs even in species, in which females lay eggs in early stages of embryogenesis. Thermal regimes simulating body temperatures of reproductive and non-reproductive females had a non-significant influence on egg size, developmental rates, and hatchling size. These results suggest that thermoregulatory behavior in females evolved earlier than advanced reproductive modes, and the phenomenon is not explainable by maternal manipulation of offspring phenotypes. It seems likely that reproductive females modify thermoregulatory behavior to maximize their egg-laying rates. Poděkování Na tomto místě bych chtěla velmi poděkovat Mgr. Lumíru Gvoždíkovi, Ph. D. za uvedení do problematiky, četné rady a připomínky a také za trpělivost a vstřícný přístup. Práce byla financována grantem Grantové agentury České republiky (P506/10/2170). Správa CHKO „Žďárské vrchy“ pro účely tohoto projektu vydala výjimku z ochranných podmínek čolka horského (1154/ZV/2008). Prohlášení Prohlašuji, že jsem svoji diplomovou práci vypracovala samostatně s využitím informačních zdrojů, které jsou v práci citovány. Brno 10. května 2013 ……………………………… Eliška Toufarová Obsah Obsah ................................................................................................................................ 9 1. ÚVOD ......................................................................................................................... 10 1.1. Mateřský efekt ...................................................................................................... 10 1.2. Vliv reprodukce na termoregulační chování ektotermních živočichů ................. 14 1.3. Vliv teploty během embryogeneze na fenotyp potomstva ................................... 18 1.4. Vliv reprodukce na termoregulační chování obojživelníků ................................. 20 1.5. Cíle práce ............................................................................................................. 22 2. MATERIÁL A METODIKA ...................................................................................... 24 2.1. Modelový organismus a jeho chov ...................................................................... 24 2.2. Měření preferovaných tělesných teplot ................................................................ 26 2.3. Měření vlivu teploty na fenotyp potomstva ......................................................... 27 2.4. Statistická analýza dat .......................................................................................... 30 3. VÝSLEDKY ............................................................................................................... 31 3.1. Termoregulační chování samic ............................................................................ 31 3.2. Fenotyp potomstva ............................................................................................... 33 4. DISKUZE ................................................................................................................... 37 4.1. Termoregulační chování samic ............................................................................ 37 4.2. Vliv samice na fenotyp potomstva ....................................................................... 39 4.3. Závěr .................................................................................................................... 40 5. LITERATURA ........................................................................................................... 42 9 1. ÚVOD 1.1. Mateřský efekt Mateřský efekt můžeme definovat jako negenetický vliv matky nebo prostředí, ve kterém se matka vyskytuje, na fenotyp potomstva (MARSHALL & ULLER 2007). Dlouho se předpokládalo, že fenotyp potomstva je pouze výsledkem interakce mezi jeho genotypem a prostředím, nicméně poznatky z evoluční biologie ukázaly, že tento předpoklad není zcela úplný (BERNARDO 1996a). V současné době je mateřský efekt považován za jeden z nejdůležitějších faktorů, který ovlivňuje fenotyp potomstva (BERNARDO 1996a; MOUSSEAU & FOX 1998). Setkáváme se s ním u většiny taxonů (MOUSSEAU & FOX 1998) a může ovlivnit téměř každý aspekt fenotypu potomků, včetně pohlaví (např. ROOSENBURG 1996), morfologie (např., DEEMING & FERGUSON 2004), a chování (např. DOWNES & SHINE 1999). Při definování mateřského účinku je důležité zdůraznit, že tento typ působení na fenotyp potomstva není omezen pouze na matky. Fenotypy „otců“ mohou mít také přímý vliv na fenotyp potomstva (GIESEL 1988 in BERNARDO 1996a). To je například u druhů, u kterých se o potomstvo stará otec (CLUTTON-BROCK 1991), nebo přes nutriční investice jako jsou svatební dary (např. NISBET 1973 in BERNARDO 1996a), či v případě, kdy fenotyp otce ovlivňuje nakladení vajec- buď obranou území, nebo budování hnízda (HOWARD 1978). Pro tyto případy lze použít obecnější termín „rodičovský efekt“. KIRKPATRICK & LANDE (1989) rozdělují mateřský efekt do dvou kategorií: první představuje nemendelovský přenos informace z matky na potomstvo, na příklad když velikost matky ovlivňuje velikost potomků, v tomto případě se jedná o tzv. "mateřskou dědičnost". V druhém případě jde o „mateřskou selekci“, což jsou případy, kdy rodiče ovlivňují selekční tlak způsobem, který přímo nesouvisí s fenotypem potomstva, například obrana snůšky proti predátorům nebo úprava hnízdního prostředí. Mateřský efekt je někdy interpretován jako důležitý mechanismus, který umožňuje adaptivní fenotypovou odpověď na očekávané podmínky prostředí. Jinými slovy, samice mohou upravit fenotypy svých potomků v reakci na podněty prostředí, které vnímá, a to může vést ke zvýšení zdatnosti (fitness) jejího potomstva v budoucnosti – hypotéza mateřské manipulace (WEBB et al. 2006). Mateřský efekt tak může mít 10 významný vliv na vývoj celé populace (KIRKPATRICK & LANDE 1989). Některé důsledky mateřského efektu však nemusí být adaptivní, ale může se jednat spíše o vedlejší fyziologické efekty (FOX & CZESAK 2000). V dřívějších studiích byl mateřský efekt považován za adaptivní pouze v případě, že zvyšoval fitness potomstva (FOX & CZESAK 2000, MOUSSEAU et al. 2009). Nicméně z pohledu potomka může být mateřský efekt nejenom adaptivní, ale také selekčně neutrální nebo dokonce negativní, když snižuje průměrnou fitness potomstva (BERNARDO 1996a, 1996b). V současné době se za hlavní měnu považuje spíše fitness matky než jejího potomstva (MARSHALL & ULLER 2007). Jednoduše by se dalo říct, že zvýšením fitness
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