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Beauveria Bassiana UNIVERSIDAD DE SALAMANCA FACULTAD DE BIOLOGÍA DEPARTAMENTO DE MICROBIOLOGÍA Y GENÉTICA MICOVIRUS ASOCIADOS A LOS HONGOS ENDOFÍTICOS Y ENTOMOPATÓGENOS Tolypocladium cylindrosporum y Beauveria bassiana Noemí Herrero Asensio 2011 UNIVERSIDAD DE SALAMANCA FACULTAD DE BIOLOGÍA DEPARTAMENTO DE MICROBIOLOGÍA Y GENÉTICA MICOVIRUS ASOCIADOS A LOS HONGOS ENDOFÍTICOS Y ENTOMOPATÓGENOS Tolypocladium cylindrosporum y Beauveria bassiana Memoria presentada por NOEMÍ HERRERO ASENSIO para optar al grado de DOCTORA EN BIOLOGÍA Salamanca, a __ de marzo de 2011 DR. ÍÑIGO ZABALGOGEAZCOA GONZÁLEZ, INVESTIGADOR CIENTÍFICO DEL CONSEJO SUPERIOR DE INVESTIGACIONES CIENTÍFICAS (CSIC), EN EL INSTITUTO DE RECURSOS NATURALES Y AGROBIOLOGÍA DE SALAMANCA, CERTIFICA Que la memoria titulada “MICOVIRUS ASOCIADOS A LOS HONGOS ENDOFÍTICOS Y ENTOMOPATÓGENOS Tolypocladium cylindrosporum y Beauveria bassiana”, presentada por Dña. Noemí Herrero Asensio para optar al grado de Doctora en Biología por la Universidad de Salamanca, ha sido realizada bajo mi dirección, en el Departamento de Estrés Abiótico del Instituto de Recursos Naturales y Agrobiología de Salamanca del Consejo Superior de Investigaciones Científicas (CSIC). Y para autorizar su presentación y evaluación por el tribunal correspondiente, expide y firma el presente certificado en Salamanca, a __ de marzo de 2011. Fdo. Dr. Iñigo Zabalgogeazcoa González DR. JOSÉ MARÍA DÍAZ MÍNGUEZ, PROFESOR TITULAR DEL DEPARTAMENTO DE MICROBIOLOGÍA Y GENÉTICA DE LA UNIVERSIDAD DE SALAMANCA, CERTIFICA Que la memoria titulada “MICOVIRUS ASOCIADOS A LOS HONGOS ENDOFÍTICOS Y ENTOMOPATÓGENOS Tolypocladium cylindrosporum y Beauveria bassiana”, presentada por Dña. Noemí Herrero Asensio para optar al grado de Doctora en Biología por la Universidad de Salamanca, ha sido realizada bajo la dirección del Dr. Iñigo Zabalgogeazcoa González, en el Departamento de Estrés Abiótico del Instituto de Recursos Naturales y Agrobiología de Salamanca del Consejo Superior de Investigaciones Científicas (CSIC), y bajo mi tutela. Y para autorizar su presentación y evaluación por el tribunal correspondiente, expide y firma el presente certificado en Salamanca, a __ de marzo de 2011. Fdo. Dr. José María Díaz Mínguez DR. ÁNGEL DOMÍNGUEZ OLAVARRI, DIRECTOR DEL DEPARTAMENTO DE MICROBIOLOGÍA Y GENÉTICA DE LA UNIVERSIDAD DE SALAMANCA, CERTIFICA Que la memoria titulada “MICOVIRUS ASOCIADOS A LOS HONGOS ENDOFÍTICOS Y ENTOMOPATÓGENOS Tolypocladium cylindrosporum y Beauveria bassiana”, presentada por Dña. Noemí Herrero Asensio para optar al grado de Doctora en Biología por la Universidad de Salamanca, ha sido realizada bajo la dirección del Dr. Íñigo Zabalgogeazcoa González, en el Instituto de Recursos Naturales y Agrobiología de Salamanca, del Consejo Superior de Investigaciones Científicas (CSIC), y tutelada por el Dr. José María Díaz Mínguez, profesor titular del Departamento de Microbiología y Genética de la Universidad de Salamanca. Y para que así conste, expido y firmo el presente certificado en Salamanca, a __ de marzo de 2011. Fdo. Dr. Ángel Domínguez Olavarri Quisiera dar las gracias a todas aquellas personas que, de algún modo, han participado en la realización de esta Tesis Doctoral: A mi director de tesis, Iñigo Zabalgogeazcoa, por confiar tanto en mí y darme la oportunidad de comprobar que la investigación, pese a todas las trabas del sistema, es un mundo apasionante y en el que quisiera quedarme. Gracias por dejarme siempre llevar a cabo mis iniciativas, por tus consejos y por apoyarme tanto, sobre todo en esta recta final que está siendo toda una locura… Al Dr. Robert Coutts por todos los consejos y enseñanzas recibidas, pues han sido de gran ayuda en la ejecución de esta tesis. A mis compañeros del grupo de pastos: Balbino, Toñi, Carlos, Beatriz, Luis, Amador, Virginia, y Cristina. Ha sido todo un placer compartir con vosotros estos cuatro años que tanto han significado para mi, tanto en lo profesional como en lo personal. Muchas gracias a todos por ser tan buenos compañeros y ayudarme tanto. Beatriz, mil gracias por esta portada de tesis tan chula!, eres una artista. A mis compañeras de labo: Lorena, que grandes momentos compartimos las tres….esos jueves tarde…jajajaj!; Encarna, muchas gracias por todos los consejos que me has dado y por ser tan buena compañera; Salud, porque además de ser una buenísima compañera, te has convertido en una persona súper importante para mi, gracias por estar siempre ahí y por ayudarme a afrontar siempre mis problemas. A mis compis del café, por ser la alegría de la huerta y hacer que las mañanas de trabajo tengan un toque diferente. Muchas gracias a probeta y precipitado, (ó Bob y Patricio? jaja), no, en serio, Sergio, Jorge, gracias por ser capaces de sacarme una carcajada a la menor y por darme tan buenos consejos siempre. A Luis, por ser el bibliotecario más enrollado que se puede tener. Y a Yolanda, por aguantarnos siempre con una sonrisa. A Silvia y Aitor, porque además de buenos compañeros de andadas en mis inicios en el IRNA, os convertisteis en unos grandes amigos. Gracias por todo! A todos los compañeros que he conocido y me han ayudado durante estos cuatro años de tesis en el IRNA. A mis compis de labo del Imperial y a mis flatmates, por hacer que recuerde mi estancia en Londres con grandísimo cariño. Al Dr. José María Díaz Mínguez, porque además de ser uno de mis mejores profes en la Uni, tuteló esta tesis doctoral. Al Dr. Ricardo Pérez y la Dra. Ana Oleaga, por introducirnos en el mundo de las garrapatas. Al Dr. Enrique Quesada Moraga, por su ayuda y por brindarnos esa impresionante colección de Beauverias. A mi familia: Mil gracias a mis padres y a mi hermana por aguantarme siempre en todos los altibajos por los que he pasado durante esta tesis. Sin vuestro apoyo incondicional esto no hubiera llegado a puerto, sois los mejores! A mi abuela Carmina, por ser tan cariñosa y darme siempre su apoyo, y a mi abuelo David, porque sé que te hubiera encantado compartir conmigo este momento. A mis tíos y primos por estar siempre pendientes de mí y ayudarme siempre a que todo fuese más llevadero. A todos mis amigos, los de siempre, y a todos los que he hecho durante estos cuatro años. Gracias por estar siempre animándome en los momentos de bajón, y sobre todo por dar el toque festivo a esta tesis, sin esas buenas fiestas que nos hemos corrido esto no hubiera sido lo mismo. Esta tesis ha sido financiada con una beca I3P CSIC y el proyecto AGL2008-01159 AGR: “Mutualismo entre hongos endofíticos y gramíneas: el papel de Epichloë festucae y sus mycovirus en la adaptación de Festuca rubra a situaciones de estrés”. Ministerio de Ciencia e Innovación + FEDER. ÍNDICE Página 1. INTRODUCCIÓN GENERAL …………………………………………. 1 1.1. Consideraciones generales sobre los hongos endofíticos…………… 3 1.2. Hongos endofíticos y entomopatógenos, dos grupos aparentemente unidos………………………………………………...……………… 5 2.1. Tolypocladium cylindrosporum …………………………………. 6 2.2. Beauveria bassiana ……………………………………………... 8 1.3. Características generales de los micovirus y su paralelismo con los hongos endofíticos ……............................................................................... 11 2. METODOLOGÍA, OBJETIVOS, RESULTADOS Y DISCUSIÓN…….. 17 Capítulo I. Abundancia de micovirus en diferentes especies de hongos endofíticos de gramíneas ……………………………………….............. 21 Capítulo II. Patogenicidad contra garrapatas, termotolerancia e infección por virus en Tolypocladium cylindrosporum…………………. 31 Capítulo III. Micovirus que infectan al hongo endofítico y entomopatógeno Tolypocladium cylindrosporum………………………. 55 Capítulo IV. Capacidad endofítica del hongo entomopatógeno Tolypocladium cylindrosporum en plantas de tomate y judía……………………………………………………………………... 81 Capítulo V. Caracterización molecular y distribución de micovirus en el hongo entomopatógeno Beauveria bassiana en España y Portugal……............................................................................................. 91 3. DISCUSIÓN GENERAL…………………………………………………. 117 3.1. Incidencia y diversidad de micovirus en hongos endofíticos y entomopatógenos……………………………………………………….. 119 3.2. TcV1 y BbRV1, dos nuevos victorivirus que infectan hongos entomopatógenos ……………………………………………………….. 122 3.3 Naturaleza dual de T. cylindrosporum: acaricida y endofito………... 126 3.4. Efecto de los micovirus en T. cylindrosporum……………………... 127 4. CONCLUSIONES ……………………………………………………….. 131 5. APÉNDICE ……………………………………………………………… 135 6. BIBLIOGRAFÍA ………………………………………………………... 153 1. INTRODUCCIÓN GENERAL Introducción general 1.1. Consideraciones generales sobre los hongos endofíticos. Los hongos endofíticos habitan en el interior de las plantas durante todo o parte de su ciclo de vida sin causarles síntomas aparentes. Se conoce de la existencia de este tipo de hongos desde finales del siglo XIX, cuando Guerin (1898) y Vogl (1898) descubrieron hongos en el interior de semillas de cizaña (Lolium temulentum). No obstante, estos hongos no recibieron especial atención hasta bien entrado el siglo XX. Fue precisamente en 1977 que se impulsó el estudio de los endofitos, cuando Charles Bacon y sus colaboradores demostraron que las toxicosis observadas en ganado vacuno estadounidense, eran debidas al consumo de plantas de Lolium arundinaceum (=Festuca arundinacea), cuyas hojas y tallos estaban infectados sistémicamente por el hongo Neotyphodium coenophialum, sin que las plantas mostrasen ningún síntoma (Bacon et al., 1977). Más tarde se descubrió que las toxicosis eran debidas a la producción de alcaloides por parte de N. coenophialum. Era
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