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Tesis Doctoral PROGRAMA DE DOCTORADO EN INVESTIGACIÓN BIOMÉDICA TESIS DOCTORAL: ENGINEERING RESPONSIVE AND BIOMIMETIC MATERIAL BASED ON ELASTIN-LIKE RECOMBINAMERS FOR BIOMEDICAL APPLICATION Presentada por Filippo Cipriani para optar al grado de Doctor/a por la Universidad de Valladolid Dirigida por: José Carlos Rodríguez Cabello Israel González de Torre Gracias a la vida que me ha dado tanto Me ha dado la marcha de mis pies cansados Con ellos anduve ciudades y charcos Playas y desiertos, montañas y llanos. Gracias a la vida – Violeta Parra (1966) ACKNOWLEDGMENTS - AGRADECIMIENTOS - RINGRAZIAMENTI Muchos años después, sobre un avión que conducía desde Florencia a Madrid, el suscrito había de recordar aquella tarde remota en que abrió Google Maps y busco donde estaba Valladolid. Me encuentro sobre el vuelo que va desde Florencia a Madrid y me acuerdo de la primera vez que vine a Valladolid; no sabía dónde estaba y nunca hubiera pensado de vivir casi 4 años en esta ciudad. Pero, si lo pienso bien, tengo que confesar que siempre he tenido un débil por España. No sé, será el Mediterráneo que nos acomuna y nos acerca; también será por el idioma tan sencillo de aprender que seguramente en estos agradecimiento estarán muchos errores. Poco importa, alguien demostró que el 90% de las comunicación nos es verbal y creo que con esto escrito el mensaje llegará aunque haya muchos errores gramaticales. Quiero empezar diciendo que haber tenido la oportunidad de vivir en este país ha sido una gran fortuna y tengo que agradecer muchas persona que han permitido que mi recorrido pasase por aquí. Habría muchos que agradecer, pero intentaré ser los más conciso posible para no aburrir demasiado. En primer lugar quiero agradecer mis directores Carlos y Israel. Muchas gracias por haberme dado esta oportunidad y por haberme guiado a lo largo del camino hasta conseguir el título de Doctor en Investigación Biomédica. Muchas gracias Carlos por tus preciosos consejos, tu enorme conocimiento sobre la ciencia, por tu generosidad y para haberme dado la libertad necesaria para desarrollar recursos que si no nunca habría descubierto. Muchas gracias Israel por tu inmenso ayudo, tu capacidad de aclararme las idea, de animarme en los momentos bajos y por haberme reconducido a las cosas prácticas cuando me estabas perdiendo en mil vueltas mentales. Muchas gracias a Matilde por tu ayuda tanto científica cuando humana, toda las veces que he entrado en tu despacho en momentos críticos, siempre he salido con más fe en mi camino y en mí mismo. Muchas gracias a Javier por tu preciosos consejos sobre el mágico mondo de la Biología Molecular; gracias a Alessandra por tu consejos sobre los experimentos de células y también para los buenos ratos pasados a hablar de la nuestra Italia y de las orígenes Etruscas que tenemos. Gracias a Luis y Menchu porque trabajar con vosotros es un auténtico lujo, donde se aprende lo que es ser científico y trabar con profesionalidad. Gracias a Merche y Irene, que me han dado mucho cariño y sencillamente verlas con esa sonrisa me animaban el corazón. Gracias a Leander y Soraya, mi inseparable compañeros del despacho. Madre mía cuantas horas pasadas untos confrontándonos sobre los problemas y ayudándose para solucionarlos. Nos hemos animado muchísimo, tanto dentro como fuera del laboratorio. Muchas Gracias a Juan y Sofía por todas las risas que nos hemos hechos, por todas las frases épicas que nos han ayudado a tener un ánimo alegre también cuando buscábamos cazar gnomos y no encontrábamos ninguno. Gracias a Tatiana y Sergio por los buenos momentos pasados en muchos congresos; gracias a Ito para sus consejos sobre que canción escuchar para motivarse y sobre los rinconcitos más bonitos de Valladolid; gracias a Arturo y Doriana porque nunca olvidaré mi primer congreso con vosotros en Oporto. Gracias a Fer, Miguel y Marcos porque MAAACCHHOOO este es un cachondeo! (y otros mil dichos que nos han hechos reír mucho). Gracias a Rocío y a Sandra por vuestra ayuda. Rocío eres una crack (como se dice aquí). Siempre lista para ayudar, tanto profesionalmente como humanamente, te mereces lo mejor. Gracias a los nuevos como Diana y Sara, que por mala suerte no hemos tenido la oportunidad de pasar mucho tiempo untos, pero estoy convencido que estaremos en contacto y nos volveremos a ver. Gracias a Sandra por haberla ayudada con su proyecto, te deseo lo mejor por tu trayectoria científica. Gracias también a lo que se han marchados antes que yo: Mohammed, Lorena, Elena, Leticia, Esther, Alicia, Teresa y Dasha, gracias por los buenos momentos pasados untos. También quiero agradecer a Gastón y sus historias sobre Cuba, al final consiguió convencerme a visitarla. Gracias también a todos los colaboradores externos, como los colegas conocidos en las estancias, los médicos de Valladolid y todos los coautores de los artículos. Gracias mis fantásticos compañeros del BIOGEL, tanto los estudiantes cuanto los Seniors, compartir con vosotros este camino profesional ha sido una oportunidad estupenda: agradezco el día que decidí aplicar por este proyecto. He aprendido muchísimo desde cada uno de vosotros, en un sentido común he aprendido lo que significa hacer ciencia y ser científico. A todos vosotros os deseo lo mejor. Terminados los agradecimientos profesionales, quiero agradecer también mi mundo fuera del Doctorado, un mundo donde conseguías recargar las pilas para volver a dar lo mejor. Gracias a todos los amigos conocidos en Valladolid; amigos inolvidable con los cuales hemos creado una conexión especial; nombrarlos todos sería demasiado, por esto me limito a decir un GRACIAS general y que llegue a todos, desde los chicos de la cafetería, pasado por los compañeros de balonmano hasta a quien me cogió en Valladolid como en una Familia. Gracias también a todos los amigos fuera de Valladolid: a lo de España, de Holanda, de Alemania, a mi amigos históricos de Italia, a los nuevos que conocí en los congresos y a lo que conocí en otros rincones del mundo. Gracias a mi Familia, porque no obstante la distancia su ayudo me ha llegado intacto y poderoso. Gracias por todas las veces que habéis venido a visitarnos recordándonos cuanto nos queréis. Que cuando estaba por el mundo conociendo gente de cultura diferente, llevaba conmigo mi mundo y mi raíces y me sentía como si vosotros estuvierais conmigo. 4 años inolvidables! Hechos principalmente de momentos buenos, pero también de momentos difíciles… como el clima de Valladolid. Niebla donde no se ve nada y solo se puede proceder a tentones. Un sol que pega tan fuerte encima de la cara que solo hay que cerrar los ojos y no permitir a las gotas de sudor de entrar. Un frio que penetra los pies porque anden más rápido, de manera de calentarse y alcanzar una meta más lejana. 4 años y mucho más pasado untos! Donde cuando había niebla andabas tu delante de mí; que cuando teníamos el sol en la cara que no se veía nada, estabas tú mirando el camino; y que cuando hacia frio para pararse, me abrazabas y me decías que la meta estaba cerca. Sin ti este reto no hubiera sido posible. Gracias a ti, Sara por todos los momentos que hemos vivido… y que tenemos que vivir. Filippo INDEX ABSTRACT ......................................................................................... 22 INTRODUCTION ............................................................................. 28 PROBLEM STATEMENT AND HYPOTHESIS ...................... 38 OBJECTIVES ...................................................................................... 42 STATE OF THE ART ........................................................................ 46 CHAPTER 1: ELASTIN-LIKE POLYMERS: PROPERTIES, SYNTHESIS AND APPLICATIONS ............................................ 48 1. INTRODUCTION ........................................................................ 49 2. ELASTIN-LIKE RECOMBINAMERs ENGINEERING, BIOPRODUCTION AND DESIGN ................................................................................... 52 2.1. History and evolution of the synthesis of elastin-like recombinamers .......................................................................... 52 1.3.1. Ancient times (the “chemistry ages”) ...................... 52 2.1.2. Modern times (the “recombinant ages”) ................ 52 2.1.3. Contemporary times (the “seamless recursive ages”)……………………………………………………………………………………53 2.2. Hosts for the expression of elastin-like recombinamers……………………………………………………………………….56 2.2.1. Prokaryotic hosts ...................................................... 56 2.2.2.1 The gold standard: Escherichia coli ................... 56 2.2.2. Eukaryotic hosts ........................................................ 58 2.2.2.1 Aspergillus nidulans fungus ............................... 58 2.2.2.2 Yeast ................................................................... 58 2.2.2.3 Plants .................................................................. 59 2.3. Novel design of elastin-like recombinamers with different features ....................................................................................... 61 2.3.1. Substitution of the guest amino acid ....................... 61 2.3.2. Fusion of other protein polymers ............................ 62 2.3.3. Fusion of bioactive domains .................................... 63 2.3.4. Fusion of full-length proteins ................................... 64 3. STRUCTURES AND PHYSICAL CHARACTERISTICS OF ELASTIN-LIKE RECOMBINAMERs .......................................................................... 65 3.1. Micelles and nanoparticles .........................................
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