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Control Por Fosfato En La Biosíntesis Del Inmunosupresor Tacrolimus En Dos Especies Del Género Streptomyces

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Control Por Fosfato En La Biosíntesis Del Inmunosupresor Tacrolimus En Dos Especies Del Género Streptomyces Universidad de León Departamento de Biología molecular INBIOTEC Instituto de Biotecnología de León Control por fosfato en la biosíntesis del inmunosupresor tacrolimus en dos especies del género Streptomyces MIRIAM MARTÍNEZ CASTRO LEÓN 2011 = = = A mis padres Agradecimientos La realización de esta tesis doctoral ha sido posible gracias a la ayuda, apoyo y ánimo que he recibido de muchas personas. El mundo de la ciencia es un mundo fascinante pero en ocasiones muy duro, momentos que se quedan en el olvido gracias al cariño que he sentido por muchos de los que me rodean. A Juan Francisco Martín, Director de esta Tesis, por haber confiado en mí al darme la oportunidad de realizar este trabajo bajo su dirección. A Carlos Barreiro, Co-director de esta Tesis y compañero, por todas tus enseñanzas y consejos pero sobre todo por la paciencia en los momentos de incertidumbre y desanimo. Gracias por enseñarme a poner en todo lo que hago un cuidado exquisito. Al Ministerio de Educación por haberme otorgado una beca predoctoral, así como la financiación para la realización de estancias en el extranjero y a la Universidad de León por ofrecerme el apoyo administrativo y de diversos recursos para la consecución de este trabajo. A todos los compañeros de Inbiotec por crear un ambiente de trabajo cálido; en general, siempre he recibido buenas palabras y disposición cuando he requerido ayuda o colaboración. En especial quiero dar las gracias a todas las personas que han pasado por mi “cuarto” con las que he compartido el trabajo del día a día codo con codo. A Antonio por la paciencia y amabilidad con las que me escuchas y aconsejas cada vez que tengo dudas sobre el complejo mundo de los Streptomicetos. A Ramiro y a José, por socorrerme en los momentos de colapso informático. A Elena, compañera de viaje en algún que otro congreso y a Javi, que me procuró medio de transporte y “culturizantes” conversaciones para la vuelta a casa durante muchos fines de semana. A los compañeros de descansos Silvia, Carlos P, Pedro y otros por ayudarme a desconectar; que bien vienen esos momentos para enfrentarse de nuevo al trabajo. A Rosma y a Patricia, por esa sonrisa cada vez que les llevaba unas muestras a secuenciar. A Claudia, Susi y Maite por sus consejos y conversaciones en las campanas o por los pasillos. A Alberto, Alma, Ana, Carlos G, Carmen, Irene, Fernando T, Katarina, Mafe, María, Marta, Nuria, Ricardo, Rosma, Rubén, Saeed, Tamara, Vane-Vane y Zahra por vuestro compañerismo. To my Ljubljana mates and Petko, which made me feel part of its group and showed me another way of working with Streptomyces. A Josefina, Berna, Andrea, Bea y Alcira, gracias a su apoyo técnico y administrativo es más fácil sacar el trabajo adelante. = Existen ciertos compañeros más especiales que no sólo me han ayudado a crecer como científica sino también como persona y con los que he tenido el placer de compartir el tiempo tanto dentro como fuera del laboratorio. Además, me han ayudado a descubrir que la ciencia no sólo se hace en el laboratorio sino que, a veces, las ideas más importantes aparecen a lo largo de interesantes conversaciones en los viajes, cenas o a la luz de las farolas. Thank you Sedef for your kind teachings and advise in my first steps in the Streptomyces world. A Mónica y a Carlos, por hacerme sentir como en casa desde el primer día, pero sobre todo por ese saber escuchar y la compañía en los malos momentos. Nunca se me olvidará ese viaje a Valencia. A Inma, Tania, Raúl y Kristian, por todos los momentos que hemos compartido. A pesar de que las circunstancias de la vida nos han deparado destinos diferentes espero que los lazos de amistad, que se forjaron aquí en León, duren para siempre. A Lorena y Rebe, mis “pequeñas” compañeras del último año, gracias por vuestro cariño y por hacerme olvidar de vez en cuando “que tenía un tesis que escribir”. A Fernando, por estar siempre dispuesto a que te bombardeara con miles de preguntas del regulón pho. A Alcira, por la simpatía y el humor desbordantes con los que has sabido ganarte a esta burgalesa. A mi grupo de “cenitas” Abel, Chi, Pili, Rubén y Ruti, por aportarme los momentos de desconexión del mundo científico y otros puntos de vista de ver la vida. Gracias por aceptar que en algunas ocasiones no podía estar porque seguía “encerrada” en el laboratorio. Y por supuesto, quiero agradecer el apoyo incondicional de mi familia que aunque no entendiesen muy bien que en qué consistía mi trabajo siempre me han proporcionado el aliento y cariño necesarios para seguir adelante. Gracias por estar siempre a mi lado. A Javi, contar contigo me hace más fuerte para afrontar cualquier reto de la vida. Gracias por tu confianza y ayuda. Abreviaturas ADNt ADN total o genómico E: unidad de repetición extensión de la ADNasa: desoxirribonucleasa unión Ácidoetilendiaminotetraacético antibióticoR: resistente a un determinado EDTA: antibiótico EU: unidad de extensión inestable de los antibioticoS: sensible a un determinado operadores de PhoP antibiótico ES: unidad de extensión de soporte de los Apr: apramicina operadores de PhoP ARNasa: ribonucleasa GST: glutatión-S-transferasa ARNr: ARN riobosomal HPLC Cromatografía líquida de alta resolución (High Performance Liquid ARNt: ARN de transferencia Chromatography). ARS: Agricultural Research Service kb: kilobase ATCC: American Type Culture Collection Kn: kanamicina AU: unidades de absorbancia M: molar. BSA: seroalbúmina bovina MOPS: ácido 3-(N- C: unidad de repetición que forma el morfolino)propanesulfonico] núcleo de la unión MTBE: metilterbutileter CIA: Cloroformo-isoamílico Mezclar 24 nm: nanómetro partes de cloroformo y 1 de alcohol isoamílico. nt: nucleótido CoA: coenzima A. pb: pares de bases Da: dalton. Pi: fosfato inorgánico ddNTP: didesoxinucleótido trifosfato. rpm: revoluciones por minuto DE: dominio efector de un regulador SDS: docedil sulfato sódico DMSO: dimetilsulfóxido TFA: ácido trifluoro acético dNTPs: desoxinucleótidos trifosfato. Tris: tris-hidroximetil-aminometano. U: unidades DO600: densidad óptica a 600 nm de longitud de onda UFC: unidades formadoras de colonias DR: dominio receptor de un regulador UV: luz ultravioleta DRu: unidad de repetición directa (direct WT: fenotipo salvaje (wild type). repeat unit) DSMZ: Deutsche Sammlung von Mikroorganismen und Zellkulturen GmbH DTT: ditiotreitol Control por fosfato de la biosíntesis del inmunosupresor tacrolimus en dos especies del género Streptomyces 1. Introducción 1 1.1. Sistemática del género Streptomyces ............................................................... 3 1.1.1. Descubrimiento de los Actinomicetos ............................................................ 3 1.1.2. Taxonomía del género Streptomyces .............................................................. 4 1.1.3. Posición actual del género Streptomyces en el Orden de los Actinomycetales 7 1.2. Características del género Streptomyces .......................................................... 9 1.2.1. Ciclo de vida ................................................................................................. 9 1.2.2. Particularidades genéticas .......................................................................... 12 1.2.3. Metabolismo secundario y su regulación .................................................... 15 1.3. Control por fosfato en el género Streptomyces ................................................ 24 1.3.1. El sistema de dos componentes PhoR-PhoP ................................................ 25 1.3.2. PhoB/P el regulador de respuesta............................................................... 30 1.3.3. El regulón pho ............................................................................................ 36 1.4. Tacrolimus .................................................................................................... 43 1.4.1. Inmunosupresores ...................................................................................... 43 1.4.2. Mecanismo de acción de los inmunosupresores .......................................... 45 1.4.3. Biosíntesis del inmunosupresor tacrolimus ................................................ 46 1.4.4. Agrupación génica de biosíntesis de tacrolimus .......................................... 49 1.4.5. Especies del género Streptomyces productoras de inmunosupresores ......... 51 1.5. Objetivos ....................................................................................................... 53 2. Materiales y métodos 55 2.1. Microorganismos utilizados ........................................................................... 57 2.1.1. Cepas de Escherichia coli ............................................................................ 57 2.1.2. Especies pertenecientes al género Streptomyces .......................................... 58 2.1.3. Otros microorganismos ............................................................................... 58 2.1.4. Crecimiento y conservación de los microorganismos ................................... 59 2.2. Medios de cultivo ........................................................................................... 60 2.2.1. Medios de cultivo para E. coli ...................................................................... 60 2.2.2. Medios de cultivo para Streptomyces spp. ................................................... 61 2.2.3. Medios propios de los estudios de taxonomía de Streptomyces .................... 62 2.2.4. Otros medios .............................................................................................. 65
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