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Dactylis Glomerata, Holcus Lanatus, Ammophila Arenaria Y Elymus Farctus

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Dactylis Glomerata, Holcus Lanatus, Ammophila Arenaria Y Elymus Farctus UNIVERSIDAD DE SALAMANCA FACULTAD DE BIOLOGÍA DEPARTAMENTO DE MICROBIOLOGÍA Y GENÉTICA ESTUDIO DE LA MICOBIOTA ENDOFÍTICA ASOCIADA A LAS GRAMÍNEAS Dactylis glomerata, Holcus lanatus, Ammophila arenaria y Elymus farctus Mª Salud Sánchez Márquez 2009 UNIVERSIDAD DE SALAMANCA FACULTAD DE BIOLOGÍA DEPARTAMENTO DE MICROBIOLOGÍA Y GENÉTICA ESTUDIO DE LA MICOBIOTA ENDOFÍTICA ASOCIADA A LAS GRAMÍNEAS Dactylis glomerata, Holcus lanatus, Ammophila arenaria y Elymus farctus Mª Salud Sánchez Márquez 2009 UNIVERSIDAD DE SALAMANCA FACULTAD DE BIOLOGÍA DEPARTAMENTO DE MICROBIOLOGÍA Y GENÉTICA ESTUDIO DE LA MICOBIOTA ENDOFÍTICA ASOCIADA A LAS GRAMÍNEAS Dactylis glomerata, Holcus lanatus, Ammophila arenaria y Elymus farctus Memoria presentada por Mª SALUD SÁNCHEZ MÁRQUEZ para optar al grado de DOCTORA EN CIENCIAS BIOLÓGICAS Salamanca, a 20 de febrero de 2009 DR. ÍÑIGO ZABALGOGEAZCOA GONZÁLEZ, CIENTÍFICO TITULAR DEL CONSEJO SUPERIOR DE INVESTIGACIONES CIENTÍFICAS (CSIC), EN EL INSTITUTO DE RECURSOS NATURALES Y AGROBIOLOGÍA DE SALAMANCA, CERTIFICA Que la memoria titulada “ESTUDIO DE LA MICOBIOTA ENDOFÍTICA ASOCIADA A LAS GRAMÍNEAS Dactylis glomerata, Holcus lanatus, Ammophila arenaria y Elymus farctus”, presentada por Dña. Mª Salud Sánchez Márquez para optar al grado de Doctora en Ciencias Biológicas 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 20 de febrero de 2009. 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 “ESTUDIO DE LA MICOBIOTA ENDOFÍTICA ASOCIADA A LAS GRAMÍNEAS Dactylis glomerata, Holcus lanatus, Ammophila arenaria y Elymus farctus”, presentada por Dña. Mª Salud Sánchez Márquez para optar al grado de Doctora en Ciencias Biológicas 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 20 de febrero de 2009. 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 “ESTUDIO DE LA MICOBIOTA ENDOFÍTICA ASOCIADA A LAS GRAMÍNEAS Dactylis glomerata, Holcus lanatus, Ammophila arenaria y Elymus farctus”, presentada por Dña. Mª Salud Sánchez Márquez para optar al grado de Doctora en Ciencias Biológicas 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 20 de febrero de 2009. Fdo. Dr. Ángel Domínguez Olavarri Deseo expresar mi agradecimiento a todas aquellas personas que de una u otra manera han contribuido a la elaboración de este trabajo: Al Dr. Ínigo Zabalgogeazcoa González, director de esta tesis, por darme la oportunidad de entrar a formar parte de tu grupo de trabajo y confiar en mi para el desarrollo de tu proyecto de endofitos. Gracias por el apoyo que me brindaste desde que llegué, y por toda tu inestimable ayuda, conocimientos, consejos, sugerencias y aportaciones para la realización de este trabajo de investigación, así como por la preocupación que has demostrado hacia mí tanto en lo profesional como en lo personal. A mis compañeros de ‘pastos’: Al Dr. Balbino García Criado, por sus sabios consejos y enseñanzas, tanto profesionales como personales. Gracias por estar siempre pendiente de todo y preocuparte tanto. A la Dra. Antonia García Ciudad, por tu ayuda desinteresada, tus palabras de ánimo, tus buenos consejos, y por brindarme siempre el apoyo necesario en cada momento. A la Dra. Beatriz Rodríguez Vázquez de Aldana, por tu ayuda incondicional, animarme siempre y sacarme de cualquier apuro. Muchas gracias por tus consejos, aportaciones y sugerencias en este y otros trabajos. A Luis, por tu buen humor y tu apoyo. A Carlos, por tu ayuda, y por enseñarme que hay que tener paciencia en esta vida… A Cristina, por ser una buena compañera y amiga. A Virginia, por estar siempre dispuesta a ayudar y ser tan buena compañera. A mis compañeros de laboratorio, con los que tantas horas he compartido durante estos años: A Lorena, por la energía que trajiste al laboratorio, y la ayuda que me prestante siempre, que buenos ratos pasamos… A Marcos, por apoyarme y valorarme tanto. Eres un gran amigo. A María, por la ayuda recibida. A Noemí, por ayudarme tanto siempre. Gracias por todo lo que me animas, me aguantas y me apoyas. ¡Eres la mejor compañera que se puede tener, y una gran amiga!. A todos mis compañeros del IRNA, porque siempre habéis tenido un momento para mí. Especialmente a: Aitor, Alicia, Álvaro, Ángel Luis, Ángel Valverde, Diego, Elena, Eva, José Javier, Juan Carlos, María González, Marta, Miguel Tapia, Olga, Oni, Pilar Cartes, Rebeca, Rocío Bastante, Rocío Vizcaíno, Sara, Sergio González, Sergio Salazar, Silvia y Yolanda, por vuestra ayuda, apoyo, y los buenos momentos vividos durante estos años. Al Dr. Gerald F. Bills, por tus enseñanzas sobre el mundo de la micología, y por tus aportaciones y tu revisión del trabajo. Gracias por las identificaciones de algunos hongos. Al Dr. José María Díaz Mínguez, por tus consejos, sugerencias, y la revisión del trabajo. A la Dra. Cristina Pita Yáñez, por hacerme unas fotos tan bonitas de las gramíneas. A mi familia: A mis padres, por dármelo todo, y ser tan comprensivos y pacientes conmigo. No existen palabras suficientes para agradeceros todo lo que hacéis por mí. Gracias mami por las horas que has dedicado a copiarme cosas. A mi enana, por quererme tanto y aguantarme. ¡Que haría yo sin ti!. A mis tíos, especialmente a Martín, Mercedes y Pedro Antonio, por preocuparos tanto por mí y darme tan buenos consejos. Gracias Nuncy y Pedro Antonio por vuestras sugerencias y correcciones. A mis primos, especialmente a Ana, por estar siempre pendiente de mí, animarme y apoyarme tanto. A todos mis amigos, especialmente: A Elena, por tu apoyo constante, por todo lo que me animas siempre, por saber que decirme en cada momento. Por compartir conmigo tantas cosas. Pero sobre todo gracias por ser tan buena amiga. A Sonia, por ser tan buena amiga. Gracias por escucharme, aconsejarme y estar siempre ahí, compartiendo conmigo cada momento importante de mi vida. A Carmen, Chema, David Álvarez, David Pérez, Félix, Javi, Leandro, Manolo, Luismi, María Manuel, María Morales, Noemi, Nuria, Segis, Silvia y Susana. Gracias por apoyarme, escucharme y animarme siempre. Y sobre quiero agradecer y dedicar esta tesis a RAÚL. Gracias a tí he sido capaz de llegar hasta aquí, pues eres la persona que más me ayuda, me anima, me apoya y me valora. Gracias por todas las horas que me has dedicado, aún estando hasta arriba de trabajo. Gracias por tus correcciones, sugerencias, aportaciones y consejos. Pero sobre todo, gracias por quererme tanto, cuidarme y aguantarme. Eres lo mejor que me ha pasado en la vida. “Todos somos muy ignorantes. Lo que ocurre es que no todos ignoramos las mismas cosas” Albert Einstein “Cada cosa tiene su belleza, pero no todos pueden verla” Confucio ÍNDICE Página 1. INTRODUCCIÓN ……………………………………………………………….. 7 1.1. Aspectos históricos de la investigación sobre hongos endofíticos …………... 10 1.2. Epichloë y Neotyphodium, los hongos endofíticos sistémicos de gramíneas .. 12 1.2.1. Ciclos de vida de Epichloë y Neotyphodium ………………………... 13 1.2.2. Efectos beneficiosos de Epichloë y Neotyphodium …………………. 16 1.2.3. Efectos perjudiciales de los endofitos en el ganado …………………. 18 1.3. Hongos endofíticos no Epichloë …………………………………………….. 19 1.3.1. Abundancia y diversidad taxonómica ………………………………... 20 1.3.2. Especificidad de tejidos ………………………………………………. 21 1.3.3. Especificidad por el hospedador ……………………………………… 22 1.3.4. Transmisión …………………………………………………………... 23 1.3.5. Tipos de interacción planta-hongo endofítico ………………………… 24 1.3.6. Interacciones mutualistas ……………………………………………… 26 1.4. Importancia del estudio de los hongos endofíticos …………………………… 28 1.5. Técnicas moleculares aplicadas a la identificación de hongos endofíticos …. 31 1.6. Las gramíneas ……………………………………………………………….. 33 1.6.1. Características generales ……………………………………………… 34 1.6.2. Distribución …………………………………………………………… 36 1.6.3. Importancia económica ……………………………………………….. 37 1.6.4. Especies de gramíneas analizadas …………………………………….. 39 1.6.4.1. Dactylis glomerata ……………………………………………….. 39 1.6.4.2. Holcus lanatus ……………………………………………………. 41 1.6.4.3. Ammophila arenaria ……………………………………………... 42 1.6.4.4. Elymus farctus ……………………………………………………. 43 2. OBJETIVOS ………………………………………………………………………. 45 3. MATERIALES Y MÉTODOS ……………………………………………………. 49 3.1. Recolección de muestras .…………………………………………………….. 51 3.2. Aislamiento de hongos .……………………………………………………….
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