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Presentación De Powerpoint Origin and evolution of Artemia reproductive and genetic diversity Marta Maccari PhD Thesis May 2017 REPRODUCTIVE AND GENETIC DIVERSITY GENETIC AND REPRODUCTIVE ARTEMIA ORIGIN AND EVOLUTION OF OF EVOLUTION AND ORIGIN Supervisors: Marta Francisco Amat Domenech Maccari 2017 Africa Gómez Gómez Programa de Doctorado en Biodiversidad Origin and evolution of Artemia reproductive and genetic diversity TESIS DOCTORAL Marta Maccari Mayo 2017 Origin and evolution of Artemia reproductive and genetic diversity Memoria presentada por Marta Maccari para optar al título de Doctora en Biodiversidad por la Universitat de València Fdo.: Marta Maccari TESIS DOCTORAL DIRIGIDA POR Dr. Francisco Amat Doménech Profesor de Investigación Instituto de Acuicultura Torre de la Sal – CSIC Vº Bº Dra. Africa Gómez Gómez Lecturer School of Environmental Sciences University of Hull (UK) Vº Bº Esta Tesis Doctoral se ha realizado con ayuda de una beca predoctoral (JAE) concedida a Marta Maccari por el Consejo Superior de Investigaciones Científicas y cofinanciada por el Fondo Social Europeo. La realización de la presente Tesis ha sido posible gracias a la financiación económica recibida a través del Proyectos de Investigación Ministerio de Ciencia e Innovación, CGL 2008-03277/BOS-2009-13. AGRADECIMIENTOS Este apartado podría hacerse interminable. Muchas han sido las personas que durante estos años me han acompañado en este recorrido que ha significado, sí, la realización de esta tesis doctoral, pero también un camino personal intenso, difícil, emocionante, enriquecedor, inolvidable. A través de estas líneas quisiera expresar mi más sincera gratitud a todas aquellas personas que han sido imprescindibles durante todo este tiempo. En primer lugar, quisiera agradecer a mis directores de tesis. Francisco Amat, por depositar su confianza en mí para la realización de este trabajo; por su entusiasmo, su experta dirección e incansable dedicación. África Gómez, por la ilusión que derrocha trabajando, por ser la primera que creyó que este día podía llegar; por ser fuente continua de ideas, por escuchar las mías y ayudarme a ordenarlas; por su apoyo incondicional. De vosotros he aprendido muchísimo. Gracias por la paciencia que habéis demostrado conmigo, por vuestro tiempo y esfuerzo. También quiero agradecer a Francisco Hontoria, a Juan Carlos e Inma por su disponibilidad, amistad y afecto y, por supuesto, al haber hecho mucho más ameno el trabajo. A todos los becarios del IATS, con los que he compartido muchas risas y momentos inolvidables: Elena, Germán, Rocío, Mohammed, Azucena, Alfonso, Gabi, Berta, Oli, Sebastián, Itziar, Raquel, Jose, Gregorio, Ana, Rafa, David, Vicky, Felipe y muchos otros…Gracias! Gracias especialmente a Diana por su ayuda en el laboratorio y su cariño, a Laura por su afecto y sentido de humor, a Majo por su amistad y complicidad, a Stella por la buena época juntas. En general quiero dar las gracias a todo el personal del IATS. A Charo por haberme acogido cada mañana con una sonrisa, a Eva por las horas amenas de V compañía, a Rosa y Emilio (y Felicidad) por haberme abierto las puertas de su casa, a Palmira por su inestimable ayuda, a Paco, Luís y Feli por su eficiente asistencia, a Silvia y María José para los buenos ratitos de charlas después de comer. Gracias también a Nieves por todo el cariño recibido durante estos anos. Las personas a las que igualmente debo un agradecimiento son aquellas a las que tuve la gran suerte de conocer en Inglaterra. A Carla, por su bonita amistad; por su apoyo y cariño en esta etapa final de la tesis y por hacer realidad sobre el papel cada formato y cada tabla imposibles. A Maria Jose por sus dosis de motivación, su ternura y sus buenos consejos. A Ana, Emilio, Mahir por las excursiones, los paseos, las charlas, las cenas y las películas compartidas. Guardo un recuerdo magnifico de vosotros! Por último debo agradecer a aquellos familiares y amigos que quizá no sepan muy bien a lo qué me he dedícado exactamente durante este tiempo, pero sin su apoyo hubiera sido imposible. Gracias a mis amigas de toda la vida Marianne, Fede y Manu, porque pensando en ellas puedo ser optimista en los momentos difíciles. Gracias por hacerme desconectar y también por entender las distancias y mis ausencias. Gracias a mis padres y a mi hermano. A ellos les debo lo que soy. Gracias por confiar en mí ciegamente, por respetar mi espacio, por preocuparos tanto y animarme a seguir siempre mí camino. Finalmente, gracias Libero por ser como eres, por estar siempre a mi lado, por tu generosidad, comprensión y paciencia. Gracias por hacerme la vida mas feliz! VI Table of contents Part I General introduction 1.- Reproductive systems. Sex v.s. asex……………………………………5 1.1.- Parthenogenesis, hybridogenesis, gynogenesis 1.2.- The paradox of sex 1.3.- Advantages of sex 1.4.- Are sexual lineages evolutionary dead ends? 1.5.- Genotypic diversity in parthenogens 2.- Parthenogenesis and its origin…………………………………………13 2.1.- Spontaneous origin 2.2.- Hybrid origin 2.3.- Infectious origin 2.4.- Contagious parthenogenesis 2.5.- Geographic parthenogenesis. Marginal habitats 2.6.- Parthenogenesis and geographic distribution 3.- Model organism: Artemia genus………………………………………24 3.1.- Morphology 3.2.- Ecology and life cycle 3.3.- Biodiversity and biogeography 3.4.- Rare males in Artemia 4.- Molecular markers to understand the evolution of parthenogenesis…………………………………………………………39 4.1.- Mitochondrial and nuclear DNA sequences 4.2.- Microsatellites 5.- Principal objectives and structure of this thesis…………………….44 Part II Articles Chapter 1.- Origin and genetic diversity of diploid parthenogenetic Artemia in Eurasia……………………………………………...51 Chapter 2.- Functional rare males in diploid parthenogenetic Artemia………………………………………………………….89 Chapter 3.- Laboratory generation of new parthenogenetic lineages supports contagious parthenogenesis in Artemia………..135 Part III General discussion and Conclusions Results and General discussion………………………………………….181 Future directions…………………………………………………………...193 Conclusions…………………………………………………………………195 Spanish summary………………………………………………..199 General bibliography………………………………………….219 Part I GENERAL INTRODUCTION General Introduction 4 General Introduction 1.- REPRODUCTIVE SYSTEMS. SEX V.S. ASEX The majority of eukaryotic species have adopted two main reproductive strategies: sexual reproduction and asexuality. Sexual reproduction, which predominates in most living organisms (Bell, 1982; De Meeûs et al., 2007; Schurko et al., 2009), is a process involving the fusion (fertilization) of two specialized reproductive cells called gametes, one from a male source and one from a female source. Both male and female gametes are produced by a special cell division process known as meiosis, which halves the number of chromosomes in each resulting sex cell. Fertilization may occur between gametes produced by a single hermaphrodite individual (selfing) or, in most cases, between gametes formed by different female and male individuals. So, in a sexual life cycle different stages alternate: diploid cellular life, meiosis, haploid cellular life, and fertilization. Meiosis and fertilization occur regularly in life cycles (Normarck et al., 2003). Asexuality is a less widespread strategy but it encompasses a variety of reproductive mechanisms (Schön et al., 2009). The term asexual reproduction sensu strictu implies the abolishment of sexes. In this case, it is considered synonymous to clonal reproduction or agametic reproduction. It occurs when an individual produces new individuals that are genetically identical to the parent at all loci in the genome, except at those sites that have experienced somatic mutations. This is, for example, the case of the fragmentation in colonial organism as reef-building corals and sponges and the case of the fission in unisexual organisms as echinoderms, turbellarian flatworms, and some polychaete and oligochaete annelid worms. Another case of clonal reproduction is 5 General Introduction represented by budding, which concerns the production of new individuals from small parts of the parent without the division of the parent individual. It is common in cnidarian (jellyfishes, hydras, corals and sea anemones), phoronids (horseshoe worms), entoprocts (goblet worms), urochordates (sea squirts) and trematodes (flukes) (De Meeûs et al., 2007). But in animal biology, we refer to asexual organisms as all those that have all dropped out of the regular meiotic (sexual) cycle (Schön et al., 2009). So, asexual reproduction regroups other types of reproduction that are not all cases of clonal reproduction and in which gametes cell are involved. This is the case of parthenogenesis, gynogenesis and hybridogenesis (Simon et al., 2003; De Meeûs et al., 2007). 1.1.- Parthenogenesis, hybridogenesis, gynogenesis Parthenogenesis refers to the development of eggs without fertilization. An unfertilized female gamete develops into a new organism (typically female, thelytokous parthenogenesis) without the need of male gamete. Modes of parthenogenetic reproduction fall into two main categories: apomixis or automixis, based on the presence or absence of meiosis. It will be explained it in more details later (Simon et al., 2003; Schlupp, 2005). Instead, gynogenesis (also called pseudogamy) is a form of reproduction in which fertilized eggs are replaced by diploid cells from the mother. Offspring are produced from diploid oocytes that do not undergo meiosis and male haploid sperm of a related bisexual species is needed only to trigger embryo development (Simon
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