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1.3 Xanthomonas Arboricola Pv. Pruni UNIVERSITAT POLITÈCNICA DE VALÈNCIA Escuela Técnica Superior de Ingeniería Agronómica y del Medio Natural Programa de doctorado en Recursos y Tecnologías Agrícolas INSTITUTO VALENCIANO DE INVESTIGACIONES AGRARIAS Centro de Protección Vegetal y Biotecnología Tesis doctoral Especies de Xanthomonas causantes de enfermedades en frutales de hueso y almendro: diagnóstico, diversidad genética y taxonomía PABLO LÓPEZ SORIANO Directoras: Dra. María Milagros López Dra. Ester Marco Noales Tutor: Dr. Josep Armengol Fortí Valencia, mayo de 2017 I II Dra. María Milagros López González, Profesora de Investigación del Instituto Valenciano de Investigaciones Agrarias, Dra. Ester Marco Noales, colaboradora científica adjunta del Instituto Valenciano de Investigaciones Agrarias y Dr. Josep Armengol Fortí, Catedrático del Departamento de Ecosistemas Agroforestales de la Universitat Politècnica de València CERTIFICAN: Que D. Pablo López Soriano ha realizado bajo su dirección en el Laboratorio de Bacteriología del Centro de Protección Vegetal y Biotecnología del Instituto Valenciano de Investigaciones Agrarias, el trabajo que con el título “Especies de Xanthomonas causantes de enfermedades en frutales de hueso y almendro: diagnóstico, diversidad genética y taxonomía”, presenta para optar al grado de Doctor. Para que así conste a los efectos oportunos, firman el presente certificado en Valencia, a 15 de mayo de 2017. Dra. María M. López González Dra. Ester Marco Noales Dr. Josep Armengol Fortí III IV Agradecimientos En primer lugar, deseo expresar mi agradecimiento a mis dos directoras, por aceptar la dirección de esta tesis y por haberme dado la oportunidad de formar parte del Laboratorio de Bacteriología del IVIA. A María Milagros López, Chiqui, muchas gracias por haber confiado en mí desde el primer momento, por tu gran profesionalidad y enorme dedicación a esta tesis, y por haberme transmitido una pequeña parte de tu amplio conocimiento científico, que ya es mucho. A Ester Marco, muchas gracias por tu enorme esfuerzo y apoyo todos estos años, por tus sabios consejos y por transmitirme tu alegría y serenidad incluso en los momentos más difíciles, que ha sido de gran ayuda. Gracias a Josep Armengol por haber aceptado ser mi tutor en la Universidad y por su gran apoyo y ayuda durante todos estos años. Con él empecé en el mundo de la patología vegetal y en gran parte gracias a él tomé la decisión de realizar la tesis. A todos los compañeros del laboratorio de Bacteriología del IVIA con los que he coincidido estos años, Clara, Javi, Adela, Inma, Carmina, Mari, Tere, Belén y Ramón, porque ha sido un lujo trabajar a vuestro lado. Muchas gracias por habérmelo puesto tan fácil desde el primer día, por vuestra agradable compañía y por echarme una mano siempre que lo he necesitado. Ha sido un privilegio formar parte de este laboratorio. A los vecinos del laboratorio de Virología, a Mariano por su gran alegría y optimismo y por preocuparse siempre por mí, a Mari Carmen por mostrarme siempre su apoyo y especialmente al gran Edson Bertolini, porque siempre ha estado dispuesto a ayudarme, por sus buenos consejos y su amistad. I A Rosa Peiró, por su energía, su paciencia y por ofrecerme su ayuda desinteresada en todo momento. A todos los becarios con los que he coincidido en el IVIA, Silvia, Bego, Dani, Gabi, Josep, Marta, Vidal, Ana, Félix, José F., Joaquín, Elena y Khaled, por su amistad, apoyo, comprensión, por todas las risas y todos los buenos ratos que hemos compartido tanto dentro como fuera del laboratorio. Nadie mejor que nosotros sabe por lo que hemos pasado. A Jaime Cubero, por permitirme hacer pequeñas estancias en su laboratorio del INIA en Madrid y a todo su equipo, que me acogieron y me hicieron sentir como uno más. Quiero agradecer especialmente a Jerson Garita, por su amistad, su paciencia y su inestimable ayuda en todo momento. A Ana Palacio, por su gran ayuda y aportación a este trabajo, desde el envío de numerosas cepas y material vegetal, hasta sus rápidas y minuciosas correcciones que han hecho posible que esta tesis salga adelante. A Miguel Cambra, posiblemente la persona que más sepa de X. arboricola pv. pruni de España, por todo el material que nos ha pasado, como su impresionante material fotográfico, por sus consejos y por estar siempre dispuesto a echar una mano. A Olivier Pruvost y Christian Vernière, por permitirme hacer dos estancias en su laboratorio del CIRAD en Isla Reunión, por haberme enseñado tanto durante esos meses, pero también por su simpatía, su amabilidad y por hacerme sentir como en casa estando tan lejos de ella. A todo el personal y técnicos del CIRAD, que me facilitaron muchísimo el trabajo, sobre todo Karine. Y a todos los amigos que hice allí, Borja, Lensa, Maeva, Isabelle, Morguen, Noura y especialmente Gilles y Jeremy, que me acogieron en su casa sin apenas conocerme. A todos ellos II muchísimas gracias por hacerme vivir una de las mejores experiencias de mi vida en esa maravillosa isla. A mis amigos de Rocafort, que sois como mis hermanos, muchísimas gracias por todas las quedadas, todas las cenas, todos los partidos, todos los viajes y por todos vuestros ánimos y ayuda durante todo este tiempo. Es imposible no pasárselo bien con vosotros. A mis amigos de Guadalaviar, porque aunque nos veamos menos de lo que me gustaría, habéis sido un apoyo muy importante durante estos años. Las quedadas con vosotros en el pueblo son uno de los mejores momentos del año. También quería agradecer a mis amigos de la universidad por sus constantes muestras de apoyo. A mi familia, mi madre Gloria y mis hermanos María José, Reyes y Fernando, porque sois mis pilares, es imposible expresar con palabras todo mi agradecimiento, por haber confiado siempre en mí, por todo vuestro apoyo, vuestro cariño, vuestros sabios consejos y, en definitiva, por estar siempre ahí. A mis cuñados Santi, Manolo y Raquel, y a mis sobrinos Alejandro, Raúl, Elisa, Mario, Carla, Sofía y Raquel, que son la alegría de la familia. A Neus, que apareció en mi vida durante el transcurso de esta tesis, muchas gracias por tu apoyo constante, por tu comprensión y por estar a mi lado de forma incondicional. Gracias a ti este trayecto ha sido mucho más fácil de recorrer. III IV Índice Resumen ................................................................................. IX Resum ................................................................................... XIII Abstract .................................................................................XV Capítulo 1: Introducción general .............................................. 1 1.1 El género Xanthomonas................................................................ 3 1.1.1 Descripción general ......................................................................... 4 1.1.2 Clasificación taxonómica ................................................................. 5 1.2 Xanthomonas arboricola ............................................................ 10 1.2.1 Descripción general .......................................................................10 1.2.2 Diversidad genética de X. arboricola .............................................11 1.2.3 Principales enfermedades asociadas a X. arboricola .....................13 1.3 Xanthomonas arboricola pv. pruni .............................................. 14 1.3.1 Descripción de X. arboricola pv. pruni ...........................................15 1.3.2 Distribución geográfica de X. arboricola pv. pruni .........................17 1.3.3 Impacto económico causado por X. arboricola pv. pruni ..............20 1.3.4 Gama de huéspedes de X. arboricola pv. pruni .............................21 1.3.5 Sintomatología asociada a la mancha bacteriana .........................25 1.3.6 Diagnóstico, detección e identificación de X. arboricola pv. pruni31 1.3.7 Epidemiología de la mancha bacteriana ........................................45 1.3.8 Diversidad genética de X. arboricola pv. pruni ..............................52 1.3.9 Control de X. arboricola pv. pruni ..................................................55 V Capítulo 2: Justificación y objetivos ........................................ 63 Capítulo 3: Inmunoensayo de flujo lateral para la detección in situ de X. arboricola pv. pruni en muestras de campo sintomáticas .......................................................................... 67 3.1 Abstract ..................................................................................... 70 3.2 Introduction ............................................................................... 71 3.3 Material and methods ................................................................ 74 3.3.1 Bacterial strains and growth conditions ........................................74 3.3.2 Production and characterization of the antisera against X. arboricola pv. pruni ........................................................................75 3.3.3 Lateral flow immunoassay .............................................................76 3.3.4 Specificity assays ............................................................................78 3.3.5 Sensitivity assays ............................................................................79 3.3.6 Comparison of LFIA with plate isolation and real time PCR for detection of X. arboricola pv. pruni in naturally infected samples80 3.4 Results ....................................................................................... 82 3.4.1 Specificity assays ............................................................................82
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