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Octopus Vulgaris Cuvier, 1797) (Octopus Vulgaris Cuvier, 1797)

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Octopus Vulgaris Cuvier, 1797) (Octopus Vulgaris Cuvier, 1797) DEPARTAMENTO DE ECOLOGÍA Y BIOLOGÍA ANIMAL UNIVERSIDAD DE VIGO UNIVERSIDAD DE VIGO DEPARTAMENTO DE ECOLOGÍA Y BIOLOGIA ANIMAL COCCIDIOSIS AND MOLECULAR BASIS OF THE DEPARTAMENTO DE ECOLOGÍA Y BIOLOGIA ANIMAL IMMUNE RESPONSE OF COMMON OCTOPUS COCCIDIOSIS AND MOLECULAR BASIS OF THE IMMUNECOCCIDIOSIS(Octopus RESPONSE vulgarisAND MOLECULAR OF COMMONCuvier, BASIS 1797) OCTOPUS OF THE IMMUNE RESPONSE OF COMMON OCTOPUS (Octopus vulgaris Cuvier, 1797) (Octopus vulgaris Cuvier, 1797) MEMORIA PRESENTADA POR SHEILA CASTELLANOS MARTÍNEZ PARAMEMORIA OPTAR AL PRESENTADA GRADO DE DOCTOR POR POR SHEILALA CASTELLANOS UNIVERSIDAD DE MARTÍNEZVIGO PARA OPTARTESIS AL DO GRADOCTORAL DE DOCTOR PORVigo, España, 2013 LA UNIVERSIDAD DE VIGO SHEILA CASTELLANOS MARTÍNEZVigo, España, 2013 Vigo, España, 2013 UNIVERSIDAD DE VIGO DEPARTAMENTO DE ECOLOGÍAY BIOLOGÍA ANIMAL COCCIDIOSIS AND MOLECULAR BASIS OF THE IMMUNE RESPONSE OF COMMON OCTOPUS (Octopus vulgaris Cuvier, 1797) MEMORIA PRESENTADA POR SHEILA CASTELLANOS MARTÍNEZ PARA OPTAR AL GRADO DE DOCTOR POR LA UNIVERSIDAD DE VIGO Vigo, España, 2013 Durante el desarrollo del presente trabajo Sheila Castellanos Martínez ha recibido una beca de estudios del Consejo Nacional de Ciencia y Tecnología (CONACyT) del gobierno de México. La presente tesis doctoral ha sido desarrollada en el marco del proyecto “Bases moleculares da resposta inmune do polbo comun, Octopus vulgaris” financiado por la Xunta de Galicia (10PXIB402116P) y en las instalaciones del Instituto de Investigaciones Marinas del Consejo Superior de Investigaciones Científicas (IIM-CSIC). Dña. Ma. del Camino Gestal Mateo, Doctora en Biología, científico titular del Consejo Superior de Investigaciones Científicas INFORMA: Que la memoria titulada “Coccidiosis and molecular basis of the immune response of common octopus (Octopus vulgaris Cuvier, 1797)” que presenta Dña. Sheila Castellanos Martínez para optar al grado de Doctor por la Universidad de Vigo, ha sido realizada bajo su dirección y considerándola concluida, autoriza su presentación a fin de que pueda ser juzgada por el tribunal correspondiente. Y para que así conste, expiden y firman el presente informe en Vigo, a 25 de Abril de 2013 Fdo. Dra. Ma. Camino Gestal Mateo D. Francisco Javier Rocha Valdés, Doctor en Biología e investigador del grupo de Zoología Marina del Departamento de Ecología y Biología Animal de la Universidad de Vigo INFORMA: Que la memoria titulada “Coccidiosis and molecular basis of the immune response of common octopus (Octopus vulgaris Cuvier, 1797)” que presenta Dña. Sheila Castellanos Martínez para optar al grado de Doctor por la Universidad de Vigo, ha sido realizada bajo su tutela y considerándola concluida, autoriza su presentación a fin de que pueda ser juzgada por el tribunal correspondiente. Y para que así conste, expiden y firman el presente informe en Vigo, a 25 de Abril de 2013 Fdo. Dr. Francisco Javier Rocha Valdés Agradecimientos En este trabajo está plasmado el esfuerzo de muchas personas que directa o indirectamente han contribuido en el desarrollo del mismo. Por tanto, cabe aquí reconocer y agradecer su valiosa colaboración. Primeramente, agradezco a mi directora, la Dra. Camino Gestal (Camino pues!, pa’ los cuates) por permitirme incursionar en este ámbito de la inmunología aplicada a los pulpos. La paciencia para enseñar y conducir a esta doctoranda durante estos 4 años ha culminado en un estudio con muchas novedades (y sólo es el principio!). La iniciación en el mundo de los parásitos de pulpos se lo debo al Dr. Eric Hochberg (Santa Barbara Museum of Natural History), quien no ha dejado de ser mi tutor, amigo, colega y proveedor de muestras hasta hoy; además de ser uno de los artífices para continuar en el camino de la parasitología. Al Dr. Francisco Rocha le agradezco haber sido mi enlace con la Universidad de Vigo. Obtener pulpos vivos para desarrollar el presente estudio no fue sencillo, pero fue subsanado gracias a la Sociedad Cooperativa Galega “Mar do Morrazo”. Los individuos cuyo peso es inferior la talla mínima de captura (1 Kg) fueron obtenidos con apoyo de la Xunta de Galicia. Entre las personas que han contribuido con su esfuerzo en el desarrollo de esta tesis está Rubén Chamorro, quien amablemente me brindó siempre su apoyo y asesoría tanto en los muestreos como en el mantenimiento de los pulpos en acuario. El Dr. Marcos Pérez Losada fue un apoyo muy importante para conseguir aclarar el dilema de la verdadera Aggregata octopiana. La invaluable asesoría de la Dra. María Prado Álvarez (la gurú del citómetro) para aplicar la citometría de flujo y su disponibilidad para responder a mis dudas posteriores han hecho posible los resultados del Capítulo 2. Igualmente, al Dr. Alejandro Romero le agradezco su asesoría y ayuda con la citometría durante mis primeras incursiones. Para el estudio de los hemocitos mediante microscopía electrónica se contó con la colaboración del Dr. Alexandre Lobo-da-Cunha y el Dr. Carlos Azevedo del Instituto de Ciencias Biomédicas Abel Salazar, de la Universidad de Porto, Portugal. Cabe mencionar que el estudio de las células del cuerpo blanco de los pulpos no habría sido posible sin la ayuda del Dr. Ángel Guerra. A la Dra. Raquel Aranguren le agradezco enormemente la disposición para orientarme en distintas técnicas empleadas en el presente estudio, su disponibilidad para resolver diversas dudas además de convertirse en una valiosa amiga. Asimismo, muchas gracias a Bego, por la ayuda para realizar los cortes histológicos de varios pulpos. El primer acercamiento a la proteómica fue gracias a la Dra. Nuria Sánchez y la Dra. María Páez de la Cadena del Laboratorio de Bioquímica de la Universidad de Vigo. Sin duda, quiero agradecer la valiosa asesoría del Dr. Ángel P. Diz, quien me enseñó a desarrollar el proyecto de proteómica con seriedad y objetividad, pero nunca aburrimiento (lo conseguimos Ángel, salió algo muy bueno!). En este apartado cabe mencionar también a Sonia Prieto (Lab. Bioquímica), quien siempre tuvo a bien resolver alguna duda y prestar su apoyo logístico en cualquier momento. Durante el desarrollo de esta tesis he contado con muy buenos compañeros y cuates de laboratorio: Rosi, Eva, Pep, Camino; y las recientes adquisiciones, Paula y Laura, con quien he compartido buenos y malos momentos. Otros buenos amigos que he hecho durante este tiempo son Rocío, Alhambra, Aldo, Miguel, Mikel, Raquel, Lourdes, Rebeca (s), Patricia, Noelia y Ma. Carmen, que han sido continuos observadores y participantes de los eventos acontecidos durante estos 4 años, dentro y fuera del instituto. Quiero hacer un reconocimiento a los chicos de Informática, que atentamente han solucionado todos los problemas con la computadora; Gracias Juan Luis por tu ayuda con la impresión de la tesis (a toda leche!). Asimismo, las chicas de laboratorio Ángeles y Cristina, siempre muy accesibles y con gran disponibilidad para ayudar a conseguir cualquier información bibliográfica. Finalmente, pero no menos importante, va mi agradecimiento a mi familia. Estos 4 años, vistos desde “El Viejo Continente” han parecido muchos más. Pero pese a todo, resisten estoicamente a las ausencias debidas a la distancia, sin faltarme nunca su apoyo. También, muchas gracias a Fernando, quien se ha convertido en uno más de los pilares importantes que me sostienen. Gracias, por acompañarme en este tiempo con todos los altibajos, en los momentos de estrés, así como aquellos de buen humor y otros no tanto. Tú has vivido tanto como yo el desarrollo de esta tesis. Gracias a todos! A mi familia CONTENTS SUMMARY i RESUMEN iii I. General introduction 1 I.1 Overview of the Octopus vulgaris fishery 2 I.2 The Octopus vulgaris aquaculture 4 I.3. Pathogens affecting wild and cultured Octopus vulgaris 8 I.3.1. Pathogens of bacterial etiology 9 I.3.2. Pathogens of protozoan etiology: the eimeriorin coccidia Aggregata spp. 10 I.4. Overview of the cephalopod immune system 13 I.4.1. Cellular factors 14 I.4.2. Phagocytosis of hemocytes 15 I.4.3. Production of Reactive oxygen species (ROS) 17 I.4.4. Production of Nitric oxide (NO) 18 I.4.5. Cellular factors as tools for assessing octopus health 19 I.4.6. Humoral factors 20 I.5. Study of the cephalopod host/pathogens interaction in the ‘omics’ era 23 I.5.1 Transcriptomics 25 I.5.2 Proteomics 27 I.6 JUSTIFICATION AND OBJECTIVES 30 Chapter 1. Molecular phylogenetic analysis of the coccidian cephalopod parasites Aggregata octopiana and Aggregata eberthi (Apicomplexa: Aggregatidae) from the NE Atlantic coast using 18S rRNA sequences 31 Abstract 32 1. Introduction 33 2. Material and Methods 35 2.1 Sampling and microscopic identification 35 2.2 Isolation and purification of the parasite 35 2.3 DNA extraction 36 2.4 DNA amplification, cloning and sequencing 36 2.5 Phylogenetic analysis 37 3. Results 38 4. Discussion 41 Chapter 2. Morphologic, cytometric and functional characterization of the common octopus (Octopus vulgaris) hemocytes 44 Abstract 45 1. Introduction 46 2. Material and methods 48 2.1 Biological material and hemolymph collection 48 2.2. Morphological characterization of O. vulgaris hemocytes 48 2.2.1 Light microscopy: fresh and fixed hemocytes 48 2.2.1.1 Cells in suspension 48 2.2.1.2 Hemolymph cell monolayer 49 2.2.2. Electron microscopy analysis 49 2.2.3. Cytochemical characterization of the white body and circulating hemocytes 50 2.2.3.1. Periodic Acid Schiff (PAS) in circulating hemocytes 50 2. 2.3.2. Arylsulphatase 50 2.2.4. Flow cytometry (FCM) analysis 50 2.3. Functional characterization of O. vulgaris hemocytes 51 2.3.1. Cell counting and hemocyte viability 51 2.3.2. Phagocytosis assay 51 2.3.3. Respiratory burst (ROS production) 52 2.3.4. Nitric oxide production (NO) 52 2.4. Statistical analyses 53 3. Results 53 3.1. Morphological and cytometric characterization of hemocytes 53 3.1.1 Light microscopy: fresh and fixed hemocytes 53 3.1.2 Electron microscopy 55 3.1.3 Cytochemistry characterization of the white body and circulating hemocytes 56 3.4 Flow cytometry 58 3.2 Functional characterization of the O.
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