Aislamiento, Caracterización Estructural Y Regulación De La Expresión De Genes Implicados En La Tolerancia Al Aluminio En Centeno

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Aislamiento, Caracterización Estructural Y Regulación De La Expresión De Genes Implicados En La Tolerancia Al Aluminio En Centeno UNIVERSIDAD COMPLUTENSE DE MADRID FACULTAD DE CIENCIAS BIOLÓGICAS Departamento de Genética AISLAMIENTO, CARACTERIZACIÓN ESTRUCTURAL Y REGULACIÓN DE LA EXPRESIÓN DE GENES IMPLICADOS EN LA TOLERANCIA AL ALUMINIO EN CENTENO MEMORIA PARA OPTAR AL GRADO DE DOCTOR PRESENTADA POR Diaa Ahmed Mohamed Abd El-Moneim Bajo la dirección de los doctores César Benito Jiménez Francisco Gallego Rodríguez Madrid, 2012 © Diaa Ahmed Mohamed Abd El-Moneim, 2012 UNIVERSIDAD COMPLUTENSE DE MADRID FACULTAD DE CIENCIAS BIOLÓGICAS DEPARTAMENTO DE GENÉTICA Aislamiento, Caracterización Estructural y Regulación de la Expresión de Genes Implicados en la Tolerancia al Aluminio en Centeno TESIS DOCTORAL DIAA AHMED MOHAMED ABD EL-MONEIM MADRID, 2012 UNIVERSIDAD COMPLUTENSE DE MADRID FACULTAD DE CIENCIAS BIOLÓGICAS DEPARTAMENTO DE GENÉTICA Aislamiento, caracterización estructural y regulación de la expresión de genes implicados en la tolerancia al aluminio en centeno Memoria que para optar al grado de Doctor en Ciencias Biológicas por la Universidad Complutense de Madrid presenta DIAA AHMED MOHAMED ABD EL-MONEIM Vº Bº Director de la Tesis Prof. Cesar Benito Jiménez Prof. Francisco Javier Gallego Rodríguez MADRID, 2012 AL ALMA DE MI PADRE, A MI MADRE, A MI QUERIDA MAYSA, A MIS NIÑAS SHAZA Y HANA, A LAS ALMAS DE LOS MÁRTIRES DE LA REVOLUCIÓN DE ENERO 2011 Ningún número de experimentos, por muchos que sean, Podrán demostrar que tenga razón. Tan solo un Experimento puede demostrar que estoy equivocado (Albert Einstein) I Hace solamente cuatro años que me embarqué en este trabajo (el mundo del aluminio) y ya estoy agradeciendo a toda la gente que me ha apoyado, ayudado, sobre todo, aguantado durante este tiempo. En fin….. ¡Cómo pasa el tiempo! Quisiera mostrar mi agradecimiento a toda esa gente que me ha apoyado durante tantos años para poder realizar mi objetivo. Seguro que me olvido de alguien, pero espero que si se da cuenta me perdone. En primer lugar dirijo mi más sincero agradecimiento al Dr. Cesar Benito y al Dr. Javier Gallego, debido a que sin su excelente dirección, la realización de esta tesis no hubiera sido posible. Gracias a Cesar Benito, mejor conocido como “Mario” no tengo palabras para describirle mi admiración y gratitud. Muchas gracias por tener tanta paciencia conmigo hasta el final, su incesante esfuerzo, apoyo, y sobre todo, por la confianza en mí durante estos años. Gracias al Dr. Javier Gallego por toda la ayuda que me ha prestado especialmente en la recta final. Igualmente los agradecimientos al Prof. Dr. Concepción Romero Martínez “Conchita”, directora del Departamento de Genética, por las facilidades concedidas para la realización de esta tesis. A los diferentes profesores del Departamento, porque siempre me han apoyado. A mis compañeros del departamento que han sido mi familia durante la mayor parte de este trabajo. Dentro de muchos otros recuerdo a (Cito A) Mónica Pradillo, Javier Varas, Cecilia Oliver, Eduardo Corredor. (Genética Molecular) Patricia Marín, Alejandra Cruz, Mari Carmen, Alberto Bellido, Beatriz Matallanas, Elisa Iniesto, Bárbara Téllez. (Cito B) Miriam González, María Cuacos. (Evolutiva) Enrique Sánchez, Diego Bersabé. Efectivamente, sin la ayuda de mis compañeros del laboratorio nada de esto hubiera sido posible. A todos los que siempre han estado dispuestos a resolver mis dudas y a darme consejos. Por todo ello debo dar las gracias a todos los que me han guiado y ensañado en algún momento a Javier Silva (con el cual todos los problemas son resueltos), Ana luisa García (siempre me decía: don´t worry everything will be OK), Naike Salvador, Roberto Contreras, José Manuel Ramos, Miriam Torres, Bea Sánchez, Bea Pérez, Elisabeth. Les digo a todos: Trabajar con vosotros ha sido y sigue siendo un placer. A mis amigos de fuera del laboratorio, que tienen paciencia y aceptan la tesis como otra carga cualquiera. A Tarek, safwat, Amr, Fouad, Haytham, Mohamed moaed, Ahmed, Asem, Nasem, Waled, Adel, Mohamed Anis, Ahmed Ibrahim, Dr/ yaser, Dr/ Amal y Dr/Mohamed . II También quiero agradecer, muy especialmente a Pilar Arnáez Aguirre por su inestimable ayuda en la escritura. GRACIAS al Programa Erasmus Mundus External Cooperation Window. por haber financiado parcialmente mi estancia en España durante todo este proceso formativo. Gracias por su solidaridad eficaz y por creer que el desarrollo de las naciones depende fundamentalmente de la educación de sus ciudadanos. De igual manera, agradezco a la SCU (Universidad Canal de Suez) y a mi País Egipto, por haber aportado la contraparte de la beca de estudios. Debo dar las gracias también a mi familia y a mis amigos por aguantar mi mal humor. Gracias a mi Padre por servirme de ejemplo. Lo más importante que he aprendido, no se encuentra en los libros, me lo has enseñado tú. A mi madre por su amor, su paciencia y por ser la mejor madre del mundo. Este trabajo mamá es fruto de tu incesante espíritu luchador por la vida de tus hijos. A mis hermanos, Basem, Shady, Bahy, Mahmud y Amany. A Maysa. Por estar a mi lado; por darme equilibrio; por su infinita confianza en mí; porque todas las dificultades han sido ligeras siempre y cuando estaba conmigo. Por su generosidad. ¡¡¡¡¡GRACIAS A TODOS!!!!! . III ÍNDICE IV Índice I-INTRODUCCIÓN 1- Tolerancia al aluminio……………………………………………...……………………..3 1.1-Importancia y alcance del problema de la toxicidad por Aluminio en suelos ácidos………………………………………..………………………………….3 1.2-Distribución global de los suelos ácidos………………………………………….4 1.3-La infertilidad de los suelos ácidos…………………………….…………………7 1.4-El aumento de la población mundial………….…………………………………8 1.5-Métodos empleados para determinar el grado de Tolerancia al Aluminio10 2-La tolerancia al Aluminio…………………………………………………………………14 2.1-Control Genético de la Tolerancia al Aluminio………………….…………….15 2.1.1-Control Genético en trigo…………………………………………………...17 2.1.2-Control Genético en Centeno………………………………………………..18 2.1.3-Control Genético en Triticale………………………………………………..19 2.1.4-Control Genético en Cebada………………………………………………..20 2.1.5-Control Genético en Avena………………………………………………….20 2.1.6-Control Genético en Maíz…………………………………………………....21 2.1.7-Control Genético en Sorgo…………………………………………………21 2.1.8-Control Genético en Arroz…………………………………………….……..21 2.1.9-Control Genético en Arabidopsis…………………………………………..22 2.1.10-Control Genético en alfalfa……………………………………………….22 2.2-Base molecular de la Tolerancia al Aluminio……………………………….22 2.3-Exclusión de Al mediante exudación de ácidos orgánicos…………….……26 2.3.1-A través de ácido málico y de proteínas transportadoras de tipo ALMT1…………………………………………………………………………..26 2.3.1.1-Trabajos llevados a cabo en trigo…………………………………..26 2.3.1.2-Trabajos realizados en Arabidopsis………………………………..32 2.3.1.3-Resultados obtenidos en centeno…………………..………………35 2.3.1.4-Resultados en otras especies……………………………….……….37 2.3.2-A través de ácido cítrico y de proteínas transportadoras de tipo AltSB y de tipo MATE…………………………………………………………...……38 V Índice 2.3.2.1-Trabajos realizados en Sorgo………………………………………..38 2.3.2.2-Resultados obtenidos en cebada………………………………….40 2.3.2.3-Resultados obtenidos en centeno, trigo, maíz y arroz……..42 2.3.3-A través de oxalato………………………………………………………..…..46 2.3.4-Metabolismo de los ácidos orgánicos………………………………..….47 3-La composición de la pared celular……………………………………………………..50 3.1-Pectin Metilesterasa (PME)……………………………………………..…………52 3.2-Relación entre PME y las respuestas frente al estrés biótico y abiótico en las plantas……………………………………………………………………………..54 3.3-Los genes ALS que codifican proteínas transportadoras de tipo ABC…..56 3.4-Exudación de aniones orgánicos por deficiencia de fósforo (P)…………….57 3.5-Mensajeros que se inducen por estrés con aluminio………………………….57 4-El centeno……………………………………………………………………………………58 4.1-Taxonomía………………………………………………………………………………58 4.2-Origen del centeno cultivado………………………………………….…………….64 4.3-Morfología y citología…………………………………………………………………65 4.4-Distribución geográfica del centeno……………………………………………….68 4.5-Importancia económica………………………………………………………..…..69 4.6-Centeno y salud………………………………………………………………………..72 4.7-Relación entre el genoma de centeno y los genomas de otras especies de Poáceas…………………………………………………………………………………..72 II-OBJETIVO……………………………………………………..……………………..……77 III-MATERIAL Y METÓDOS…………………………………………………………….79 1-Material vegetal……………………………………………………………………………80 2-Métodos……………………………………………………………………………………..82 2.1-Test Básico de Tolerancia al Aluminio………………………………………..82 2.1.1-Tinción con Eriocromo de Cianina R……………………………….……..82 2.1.2-Tinción con Hematoxilina………………………………….………………83 2.2- Test Adicionales de Tolerancia al Aluminio……………………………..….84 2.2.1-Distribución y acumulación del Al…………………………………………84 VI Índice 2.2.2-Visualización de la peroxidación lipídica……………………..………….84 2.2.3-Determinación de la muerte celular……………………………………..84 2.3-Aislamiento y purificación de DNA……………………………………………..84 2.4-Aislamiento y purificación de RNA…………………………………………….85 2.5-Valoración y cuantificación de ácidos nucleicos……………………….…….85 2.6-Amplificación de ácidos nucleicos por PCR………………………………..….85 2.7-Clonación y secuenciación………………………………………….……………..86 2.8-Análisis de secuencias……………………………………………………….……..87 2.9-Identificación de secuencias flanqueantes………………………...….……….88 2.10-Análisis de la expresión génica……………………………………………..…..88 2.10.1-Transcripción inversa en ensayos de expresión génica…………..…88 2.10.2-Exposición de plántulas al aluminio…………………………...……..89 2.10.3-Determinación de la expresión génica mediante PCR semicuantitativa……………………………………………………..……89 2.10.4-PCR cuantitativa en tiempo real mediante SYBR
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