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Gasparini, Clelia (2009) Sexual conflict in the guppy, Poecilia reticulata. [Tesi di dottorato]

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Abstract (inglese)

Sexual conflict is a widespread phenomenon in nature and it occurs over a wide range of decisions involving female-male interactions. This conflict drives the evolution of sexually antagonistic traits. Under a sexual conflict scenario, a trait that is beneficial to one sex is detrimental to the other driving the evolution of counteradaptation in a sort of ‘arm race’. In this process every evolutionary change in one sex is followed by a change in the opposite sex in order to minimize costs associated to the former. Mating frequency is one of the aspects of reproduction that could especially fuel sexual conflict and consequently sexually antagonistic coevolution because generally males have an optimal mating rate higher than females. My PhD thesis is aimed to study different aspects of sexual conflict using a small livebearing fish, Poecilia reticulata, as a model system. This species is particularly suitable for this purpose since sexual conflict is evident over mating rate, as revealed doubtless from the presence of high level of sexual harassment. I divided my researches into two main trends. First, I investigated costs and benefits of sexual conflict. Second, I focused on sexually antagonistic coevolution.
Costs and benefits of sexual conflict. Estimate costs and benefits of sexual conflict shed new light on the evolution of female resistance that is assumed to evolve to minimize the costs associated to superfluous matings. However, a recent hypothesis argues that female resistance evolves as a way to screen the more successful male and therefore operate via indirect benefits mechanism. Results from a study of the effects of sexual harassment on female’s lifetime fecundity and on offspring’s quality revealed that do not support this latter hypothesis. I demonstrated only costs of sexual harassment on offspring both in daughters and in sons. Moreover, study on heritability of males’ traits influencing reproductive success both at pre- and postcopulatory level revealed scarce sire heritability that instead is a fundamental assumption of the hypothesis. The low heritability for males’ traits may be explained by adaptive phenotypic plasticity that has been investigated for sperm velocity. This demonstrated an adjustment in sperm velocity by males according to mating opportunities.
Sexually antagonistic coevolution. My study of sexually antagonistic coevolution focuses on genitalia (male gonopodium and female oviduct). Gonopodial shape and dimension result important factors in determining the outcome of both consensual and coercive mating. Gonopodial length and shape resulted to be under sexual selection by female premating choice, with females mate more quickly with males possessing longer gonopodia with an enlarged shape. Gonopodium tip shape is revealed to be important in sperm transfer both in consensual and in forced matings. Another experiment demonstrated that females from low conflict population have minor control over sperm transfer in forced copulations by high conflict population males than relative counterparts. This result highlights the importance of male-female antagonistic coevolution in a sexual conflict scenario. An intraspecific study at population level corroborates this conclusion by providing evidence for a coevolutionary pattern between male and female concerning genitalia morphology. For this reason, genitalia might be the result of an evolutionary arm race between males and females in the struggle for control over mating rate and insemination success.

Abstract (italiano)

Il conflitto sessuale è un fenomeno ubiquitario in natura e interessa molte delle interazioni maschio-femmina portando alla evoluzione di caratteristiche sessuali antagonistiche. Per definizione, nel conflitto sessuale una caratteristica che avvantaggia un sesso comporta dei costi per l’altro sesso, per cui di conseguenza in quest’ultimo viene favorita la comparsa di controadattamenti in una sorta di ‘corsa agli armamenti’, in cui ogni cambiamento evoluzionistico che appare in un sesso è seguito da un cambiamento nell’altro sesso volto a minimizzarne i costi. La frequenza degli accoppiamenti è uno degli aspetti cruciali in cui il conflitto sessuale si manifesta e porta alla selezione sessuale antagonistica perché in generale i maschi hanno una frequenza ottimale di accoppiamenti che è maggiore di quella delle femmine. La mia tesi di dottorato ha avuto lo scopo di studiare alcuni aspetti del conflitto sessuale utilizzando Poecilia reticulata come modello di studio. Questa specie è particolarmente adatta per lo studio di queste tematiche, dal momento che è evidente la presenza del conflitto sessuale per quanto riguarda il numero di accoppiamenti, come si evince chiaramente dall’alto livello di sexual harassment (i continui tentativi di copula da parte dei maschi) subito dalle femmine. Le ricerche che ho effettuato per la mia tesi di dottorato si possono suddividere in due filoni. Primo: costi e benefici del conflitto sessuale. Secondo: coevoluzione sessuale antagonistica.
Costi e benefici del conflitto sessuale. Studiare costi e benefici legati al conflitto sessuale è importante per capire l’evoluzione della resistenza femminile che si assume si sia evoluta con lo scopo di ridurre i costi legati agli accoppiamenti superflui. Di recente tuttavia si è avanzata l’ipotesi che la resistenza evolva attraverso il meccanismo dei benefici indiretti ottenuti selezionando i maschi più persistenti. Il primo lavoro ha riguardato lo studio degli effetti del sexual harassment sulla fecondità life-time delle femmine, ma anche degli effetti sulla prole. I risultati indicano che il costo del sexual harassment è visibile sulla prole, sia nei maschi che nelle femmine. Questi risultati non supportano perciò l’ipotesi che la resistenza femminile evolva attraverso l’acquisizione di benefici indiretti. Inoltre lo studio sull’ereditabilità di tratti maschili legati al successo riproduttivo ha dimostrato che tali caratteristiche sono poco ereditabili per via paterna, presupposto essenziale per questa ipotesi. La plasticità adattativa dei caratteri ne può spiegare la scarsa ereditabilità paterna riscontrata. Ho testato questa possibilità per un tratto legato al successo riproduttivo, la velocità spermatica, a seconda delle opportunità di accoppiamento percepite dal maschio. Si è dimostrato che i maschi sono in grado di modulare la velocità spermatica, confermando perciò la possibilità di un certo grado di plasticità che può spiegare i bassi valori di ereditabilità.
Coevoluzione sessuale antagonistica. Lo studio della coevoluzione sessuale antagonistica si è concentrato sulla morfologia dei genitali, sia maschili che femminili. I risultati indicano che la morfologia dell’organo copulatore influenza il successo nel maschio sia negli accoppiamenti cooperativi che in quelli forzati. In particolare la dimensione e la forma generale dell’organo copulatore sono sessualmente selezionati attraverso la scelta femminile, e le femmine si accoppiano più velocemente con maschi dotati di gonopodi più lunghi e di forma allargata. Anche la forma della parte più distale del gonopodio risulta importante nel determinare il trasferimento degli spermi.
La coevoluzione sessuale antagonistica prevede che quando una caratteristica modellata dal conflitto sessuale differisce tra popolazioni diverse, le femmine siano più resistenti ai maschi della propria popolazione, con cui si sono coevolute, rispetto a quelli di popolazioni diverse contro cui esse non hanno gli adattamenti necessari per resistere ad accoppiamenti non voluti.
Questa previsione è stata confermata attraverso uno studio su maschi e femmine di popolazioni a diverso conflitto sessuale, in cui è stato dimostrato che femmine poste con maschi provenienti da una popolazione con un maggiore grado di conflitto sessuale hanno un minor controllo sul trasferimento degli spermi, rispetto a femmine che hanno avuto la possibilità di coevolvere con tali maschi. Infine uno studio comparativo fra popolazioni diverse indica l’esistenza di coevoluzione tra genitali maschili e femminili, che varia a seconda del grado di conflitto sessuale a cui le popolazioni sono soggette. Per cui l’evoluzione dei genitali sembra essere il risultato di una corsa agli armamenti tra maschi e femmine per il controllo della frequenza degli accoppiamenti e del successo di inseminazione.

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Tipo di EPrint:Tesi di dottorato
Relatore:Pilastro, Andrea Augusto
Dottorato (corsi e scuole):Ciclo 21 > Corsi per il 21simo ciclo > BIOLOGIA EVOLUZIONISTICA
Data di deposito della tesi:30 Gennaio 2009
Anno di Pubblicazione:Febbraio 2009
Parole chiave (italiano / inglese):Conflitto sessuale; coevoluzione sessuale antagonistica; Poecilia reticulata
Settori scientifico-disciplinari MIUR:Area 05 - Scienze biologiche > BIO/05 Zoologia
Struttura di riferimento:Dipartimenti > Dipartimento di Biologia
Codice ID:1787
Depositato il:30 Gen 2009
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