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Caracterización De Bacterias Aeromonadales Móviles Aisladas De Peces Cultivados En Uruguay

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Caracterización De Bacterias Aeromonadales Móviles Aisladas De Peces Cultivados En Uruguay UNIVERSIDAD DE LA REPÚBLICA FACULTAD DE VETERINARIA Programa de Posgrados CARACTERIZACIÓN DE BACTERIAS AEROMONADALES MÓVILES AISLADAS DE PECES CULTIVADOS EN URUGUAY Alejandro Perretta TESIS DE MAESTRÍA EN SALUD ANIMAL URUGUAY 2016 i ii iii UNIVERSIDAD DE LA REPÚBLICA FACULTAD DE VETERINARIA Programa de Posgrados CARACTERIZACIÓN DE BACTERIAS AEROMONADALES MÓVILES AISLADAS DE PECES CULTIVADOS EN URUGUAY Alejandro Perretta _________________________ _________________________ Dr. Pablo Zunino Dra. Karina Antúnez Director de Tesis Codirectora de Tesis 2016 iv INTEGRACIÓN DEL TRIBUNAL DE DEFENSA DE TESIS Claudia Piccini; MS, PhD Departamento de Microbiología Instituto de Investigaciones Biológicas "Clemente Estable"- Ministerio de Educación y Cultura - Uruguay Laura Bentancor; MS, PhD Departamento de Bacteriología y Virología Instituto de Higiene - Facultad de Medicina Universidad de la República - Uruguay Martín Fraga; MS, PhD Plataforma de Salud Animal Instituto de Investigaciones Agropecuarias - Uruguay 2016 v vi vii DEDICATORIA …para Federíco, Anaclara y Valeria viii AGRADECIMIENTOS - A Pablo Zunino y Karina Antúnez, gracias por compartir el conocimiento conmigo, abrirme las puertas del IIBCE para poder llevar a cabo este trabajando y por la paciencia y el apoyo en todo momento. - A Daniel Carnevia por el apoyo contínuo y la colaboración brindada en cada uno de mis emprendimientos. - A todo el personal docente y no docente del Instituto de Investigaciones Pesqueras de la Facultad de Veterinaria por tantos años de apoyo y colaboración. - A Belén Branchiccela y todo el personal del Departamento de Microbiología del IIBCE por el apoyo y la colaboración en todas las tareas llevadas a cabo allí. - A Rodrigo Puentes, Uruguaysito Benavides y todo el personal del Área Inmunología de la Facultad de Veterinaria por la colaboración con las taréas de biología molecular llevadas a cabo. - A Amy Carnahan, P. Srinivasan, Pol Huedo y Boris Jovanovic por la colaboración en aspectos relacionados con la identificación de aeromonadales y la evaluación de virulencia de los mismos. - A los acuaristas, productores y veterinarios que aportaron casos clínicos para ésta investigación. - A Claudia Piccini, Laura Bentancor, Martín Fraga, Leonardo Galli, Rolando Mazzoni y Daniel Carnevia por la revisión crítica y los aportes realizados al documento final de tesis. - A Sci-Hub y toda la comunidad de investigadores que trabajan por una ciencia open access. - A mi familia!!! Por ser el motorcito que me impulsa día a día. ix ÍNDICE Resumen xii Summary xiii 1. Introducción 1 1.1. Aspectos taxonómicos 1 1.2. Principales características de los aeromonadales móviles 3 1.3. Aislamiento e identificación 5 1.3.1. Identificación por métodos bioquímicos 6 1.3.2. Identificación por métodos moleculares 6 1.4. Antecedentes en la caracterización de Aeromonas spp. 7 1.4.1. Caracterización bioquímica 7 1.4.1.1. Resistencia a antibióticos 9 1.4.2. Caracterización molecular 10 1.5. La enfermedad en los peces 10 1.5.1. Signos clínicos 11 1.5.2. Patogenia 11 1.5.3. Hallazgos anatomopatológicos y paraclínicos 12 1.5.4. Epizootiología 13 1.6. Importancia para la salud humana 13 2. Hipótesis 15 3. Objetivos 15 3.1. Objetivo general 15 3.2. Objetivos específicos 15 4. Materiales y Métodos 16 4.1. Origen de las aislamientos 16 4.2. Identificación fenoespecífica y caracterización bioquímica 16 4.2.1. Actividad hemolítica 17 4.2.2. Antibiograma 17 4.3. Identificación y caracterización molecular 19 4.3.1. Búsqueda de genes vinculados a factores de virulencia 20 4.3.2. Tipificación mediante técnicas de rep-PCR 20 4.3.3. Análisis de conglomerados a partir de la tipificación de los aislamientos 21 4.3.4. Análisis filogenéticos en base a las secuencias de los genes ARNr 16S y gyrB 21 4.4. Caracterización patológica 22 4.4.1. Modelo in vivo para la evaluación de virulencia 22 4.4.1.1. Sujetos experimentales 22 4.4.1.2. Confección del inoculo y administración a los peces 23 4.4.2. Hallazgos de histopatología 24 4.5. Análisis estadísticos 24 5. Resultados 25 5.1. Origen e identificación de los aislamientos 25 5.2. Caracterización bioquímica 27 5.3. Antibiograma 28 5.4. Genes de virulencia 30 5.5. Caracterización genética 34 5.5.1. Tipificación con técnicas rep-PCR 34 5.5.1.1. Empleo de primers BOX 34 5.5.1.2. Empleo de primers ERIC 37 5.5.1.3. Empleo de primers REP 40 5.5.2. Análisis filogenético a partir de las secuencias del ARNr 16S 41 5.5.2.1. Análisis intraespecífico para las especies nacionales 41 5.5.2.2. Análisis interespecífico de los aislamientos nacionales 42 5.5.2.3. Análisis filogenético realizado entre los aislamientos nacionales y las secuencias 43 depositadas en RDP 5.5.3. Análisis filogenético a partir de las secuencias del gen gyrB 45 5.5.3.1. Análisis intraespecífico para las especies nacionales 45 x 5.5.3.2. Análisis interespecífico de los aislamientos nacionales 46 5.5.3.3. Análisis filogenético entre los aislamientos nacionales y las secuencias depositadas en 46 Aeromonas MLST Databases 5.6. Caracterización patológica 53 5.6.1. Elección de aislamientos candidatos 53 5.6.2. Cálculo de DL5096 para cada aislamiento seleccionado 54 5.6.2.1. Aislamiento H2 54 5.6.2.2. Aislamiento H11 56 5.6.2.3. Aislamiento V2 57 5.6.2.4. Aislamiento V6 59 5.6.3. Comparación entre aislamientos 59 5.6.4. Hallazgos histopatológicos en los peces infectados 60 6. Discusión 63 6.1. Origen e identificación de los aislamientos 63 6.1.1. Hallazgos epizootiológicos 65 6.2. Caracterización bioquímica 66 6.3. Antibiograma 68 6.3.1. Índice de Multirresistencia a Antibióticos (MAR) 69 6.4. Caracterización genética 70 6.4.1. Genes de virulencia 70 6.4.1.1. Aeromonas allosaccharophila 72 6.4.1.2. Aeromonas bestiarium 72 6.4.1.3. Aeromonas caviae/punctata 72 6.4.1.4. Aeromonas hydrophila 73 6.4.1.5. Aeromonas veronii 74 6.4.2. Tipificación mediante técnicas de rep-PCR 75 6.4.2.1. Tipificación de Aeromonas hydrophila 76 6.4.2.2. Tipificación de Aeromonas caviae/punctata 78 6.4.2.3. Tipificación de Aeromonas veronii 79 6.4.3. Análisis filogenéticos 79 6.4.3.1. Empleo de la secuencia del gen que codifica para el ARNr 16S 80 6.4.3.2. Empleo de la secuencia del gen gyrB 80 6.5. Caracterización patológica 83 7. Conclusiones 86 8. Bibliografía 88 Anexo 1: Composición de medios de cultivo 106 Anexo 2: Protocolo de extracción de ADN y composición de solución fijadora de Davidson 107 Anexo 3: Perfiles de identificación molecular de los aislamientos nacionales de Aeromonas 108 spp. para los genes ARNr 16S y gyrB Anexo 4: Árboles filogenéticos correspondientes al análisis de los genes ARNr 16S y gyrB de 143 los aislamientos nacionales de Aeromonas spp. xi LISTA DE CUADROS Cuadro 1. Descripción de los rasgos fenotípicos de Aeromonas spp. móviles en los que se halló 66 discrepancia entre los perfiles obtenidos de aislamientos de peces de Uruguay y la información publicada (Chong et al., 1980, Carnahan et al., 1991; Abbott et al., 1992; Abbott et al., 2003). LISTA DE FIGURAS Figura 1. Corrida electroforética correspondiente a la tipificación con primers BOX realizada 35 sobre Aeromonas hydrophila (H) y A. punctata (P) aisladas de peces cultivados en Uruguay (L: ladder 10 kb). Figura 2. Corrida electroforética correspondiente a la tipificación con primers BOX de 35 Aeromonas allosacharophila (A), A. bestiarium (B) y A. veronii (V) aisladas de peces cultivados en Uruguay (L: ladder10 kb). Figura 3. Análisis de clusters realizado sobre la corrida electroforética correspondiente a la 36 tipificación de aislamientos de Aeromonas hydrophila (H1 a H13) con primers BOX mediante empleo del software Gelcompar II®. El número en cada nodo indica el valor de bootstrap calculado sobre 500 repeticiones (p<0,05) (*: indican aislamientos procedentes de un mismo brote epizoótico). Figura 4. Análisis de clusters realizado sobre la corrida electroforética correspondiente a la 36 tipificación de aislamientos de Aeromonas veronii (V1 a V11) con primers BOX mediante empleo del software Gelcompar II®. El número en cada nodo indica el valor de bootstrap calculado sobre 500 repeticiones (p<0,05) (*, +: símbolos iguales indican aislamientos procedentes de un mismo brote epizoótico). Figura 5. Análisis de clusters realizado sobre la corrida electroforética correspondiente a la 37 tipificación de aislamientos de Aeromonas punctata (P1, P2, P4 y P5) con primers BOX mediante empleo del software Gelcompar II®. El número indica el valor de bootstrap calculado sobre 500 repeticiones (p<0,05) (*: aislamientos procedentes de un mismo brote epizoótico). Figura 6. Corrida electroforética correspondiente a la tipificación con primers ERIC realizada 37 sobre Aeromonas allosaccharophila (A), A. hydrophila (H) y A. punctata (P) aisladas de peces cultivados en Uruguay. L: ladder 10kb. Figura 7. Corrida electroforética correspondiente a la tipificación con primers ERIC y REP 38 realizada sobre Aeromonas bestiarium (B) y A. veronii (V) aisladas de peces cultivados en Uruguay. L: ladder 10kb. Figura 8. Análisis de clusters realizado sobre la corrida electroforética correspondiente a la 39 tipificación de aislamientos de Aeromonas hydrophila (H1 a H3 y H5 a H13) con primers ERIC mediante empleo del software Gelcompar II®. El número en cada nodo indica el valor de bootstrap calculado sobre 500 repeticiones (p<0,05) (*: aislamientos procedentes de un mismo brote epizoótico). Figura 9. Análisis de clusters realizado sobre la corrida electroforética correspondiente a la 39 tipificación de aislamientos de Aeromonas veronii (V1 a V11) con primers ERIC mediante empleo del software Gelcompar II®. El número en cada nodo indica el valor de bootstrap calculado sobre 500 repeticiones (p<0,05) (*, +: símbolos iguales indican aislamientos procedentes de un mismo brote epizoótico).
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