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SISTEMAS DE INMERSIÓN TEMPORAL, ELICITACIÓN Y TRANSFORMACIÓN GENÉTICA EN Digitalis Purpurea L

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SISTEMAS DE INMERSIÓN TEMPORAL, ELICITACIÓN Y TRANSFORMACIÓN GENÉTICA EN Digitalis Purpurea L INSTITUTO DE BIOTECNOLOGÍA DE LAS PLANTAS SISTEMAS DE INMERSIÓN TEMPORAL, ELICITACIÓN Y TRANSFORMACIÓN GENÉTICA EN Digitalis purpurea L. Tesis presentada en opción al grado científico de Doctor en Ciencias Agrícolas Naivy Lisbet Pérez Alonso Santa Clara 2013 UNIVERSIDAD CENTRAL “MARTA ABREU” DE LAS VILLAS INSTITUTO DE BIOTECNOLOGÍA DE LAS PLANTAS SISTEMAS DE INMERSIÓN TEMPORAL, ELICITACIÓN Y TRANSFORMACIÓN GENÉTICA EN Digitalis purpurea L. Tesis presentada en opción al grado científico de Doctor en Ciencias Agrícolas Autor: Ing. Naivy Lisbet Pérez Alonso, MSc Tutor: Inv. Tit., Ing. Elio Antonio Jiménez González, Dr. C Santa Clara 2013 CITACION CORRECTA Sistema apellido‐año Pérez Alonso, Naivy Lisbet. 2013. Sistemas de inmersión temporal, elicitación y transformación genética en Digitalis purpurea L. [Tesis presentada en opción al Grado Científico de Doctor en Ciencias Agrícolas] Santa Clara, Universidad Central “Marta Abreu” de Las Villas, Instituto de Biotecnología de las plantas. 97 p. Sistema numérico 1. Pérez Alonso, Naivy Lisbet. Sistemas de inmersión temporal, elicitación y transformación genética en Digitalis purpurea L. [Tesis presentada en opción al Grado Científico de Doctor en Ciencias Agrícolas] Santa Clara, Universidad Central “Marta Abreu” de Las Villas, Instituto de Biotecnología de las plantas, 2013. 97 p. Agradecimientos Soy de pocas palabras pero para aquellos que lo merecen va todo mi esfuerzo -A ti, amigo entrañable y tutor que hace más de 18 años, me diste la oportunidad de conocerte, aprender y amar la biotecnología. De ti recibí grandes consejos en lo profesional y en lo personal y sobre todo, recibí siempre tu apoyo. No olvides que sigo pensando que cuando sea grande quiero ser como tú - Gracias a ti, Rul, por ser también mi maestro y amigo, te quiero con la vida y te agradezco, incluso, los malos momentos que nos hacen ser mejores -Al Dr André Gerth por permitirme realizar las investigaciones en BioPlanta y a la vez agradecerle a todos los que hicieron mi estancia allí más agradable. Al Dr Gerhard Kerns, Dr Horst-Michael Nitzsche, Dr Dirk Wilken y mi querida amiga Annet Jähn un millón de gracias -A aquellos que revisaron minuciosamente cada detalle de este manuscrito y que aportaron valiosas ideas y cuyos nombres quisiera mencionar pero sería imposible, especialmente le agradezco a los Doctores Gil Enríquez y Miguel Ramos que pusieron toda su energía y entrega para hacer de este, un mejor documento. A Kosky por las horas de revisión y discusión -A Marta y Osmildo por permitirme mantener temporalmente una biblioteca personal y que no han escatimado esfuerzos para ayudarme siempre. A Yeny y Lourdes por quitarme tantas preocupaciones burocráticas -A Marisol por todo el apoyo de la subdirección de investigaciones y a la dirección del IBP en la persona de Osvaldo por facilitarme todo lo necesario para la preparación y presentación de la tesis. -A ti Migue, mi viejito por cuidar de mi y ayudarme siempre así como a todo el personal de investigación, producción y servicios que han seguido de cerca cada avance de la tesis -A los amigos de siempre, Leo, Mariela, Daimy, que aún sin saberlo me han hecho crecer y llegar al final de esta carrera. A Maricarmen y Andrés, con los que viví experiencias inolvidables en la lejana Europa y de los que recibí ayuda profesional. A otros como Diego que forman parte de aquella etapa diferente en mi vida… hazte Doctor ya y te sentirás ligero Agradecimientos -A uds amigas del alma, Alina y Elisa, con quienes he compartido momentos alegres otros tristes, pero momentos al fin y de las que he aprendido el valor de la amistad. A uds que me conocen y aceptan tal como soy les agradezco estar siempre...Ellas por aparecer antes en mi vida, pero a uds, Neyda, Gre, Anabel, Ida mil gracias por la amistad. -A tía Yoly, tía Reina, abuela Yolanda, Bellito, gracias por cuidar siempre de mi hija y de mi chino con la mayor dedicación -A mis padres que me enseñaron la grandeza de estudiar y hacer el bien, a mi hermana que es mi sostén, mis tías, primos y primas, que no me pierden de vista, a mi familia entera…. A mi sobrina Adriana, que no es la única ni la más querida, porque a todos los quiero igual, pero que el nacimiento de este proyecto me impidió estar en el suyo -A ti, mi amor y amor de todas por tu apoyo, comprensión aún cuando me recuerdas lo tarde que he finalizado este proyecto y en el cual has puesto todo tu conocimiento. Por estar a mi lado, a veces despierto otras dormido, mientras las ideas iban y venían… , a ti que me diste tu fuerza cuando me sentí desfallecer, a ti que me diste la mayor alegría de mi vida, nuestra Lia -Y por supuesto, a ti, mi Lia amada, amor de mis amores, que me has visto sacrificar horas junto a ti para dedicárselas a la tesis. A ti que has sido paciente con solo 5 años y has sabido esperar, a ti que aprenderás que en la vida hay sacrificios tan grandes como estos pero que valen la pena A todos los que están y ya no están en cuerpo y alma pero sí en mi corazón, gracias Abreviaturas Descripción completa 6-BAP 6-Bencilaminopurina ACN Acetonitrilo ADN Ácido desoxirribonucleico ADN-T Ácido desoxirribonucleico de transferencia AIA Ácido indol-3-acético ARN Ácido ribonucleico AS Acetosiringona ChP ChitoPlant® EROs Especies reactivas de oxígeno gMF Gramos de masa fresca gMS Gramos de masa seca HPLC Cromatografía Líquida de Alta Resolución (del inglés High-Performance Liquid Chromatography) KAc Acetato de potasio kb kilopares de bases MDA Malondialdehido MJ Metil jasmonato MFC Medio de cultivo de formación de callos MMb Medio de cultivo de multiplicación de brotes MS Sales inorgánicas propuestas por Murashige y Skoog (1962) pb pares de bases PCR Reacción en cadena de la polimerasa (del inglés Polimerase Chain Reaction) Rif Rifampicina RITA® Recipientes de inmersión temporal automatizados SDS Sulfato duodecilo de sodio SiP SilioPlant® SIT Sistemas de inmersión temporal Spm Espectinomicina (del inglés Spectinomycin) Str Estreptomicina (del inglés Streptomycin) SÍNTESIS Digitalis purpurea L. es una de las dos únicas especies de interés económico del género Digitalis. Su importancia radica en que constituye una de las principales fuentes de cardenólidos, fármacos irreemplazables en el tratamiento de la insuficiencia cardíaca. El presente trabajo se realizó con el objetivo de estudiar diferentes estrategias biotecnológicas en Digitalis purpurea L. que permitirán disponer de la base metodológica para el desarrollo de una tecnología para la producción in vitro de cardenólidos. Para su cumplimiento se desarrolló un método de producción de biomasa en sistemas de inmersión temporal (SIT), se determinó el efecto de elicitores en el contenido de cardenólidos y se desarrolló un protocolo para la transformación genética mediado por Agrobacterium tumefaciens. La mayor producción de biomasa por SIT (104,03 gMF y 5,74 gMS) se obtuvo cuando se inocularon 12 explantes. La frecuencia de inmersión influyó en la producción de biomasa y de cardenólidos en los brotes. En el tratamiento con inmersiones cada 4 horas se obtuvo la mayor producción de biomasa y producción neta de cardenólidos por SIT (digitoxina 167,6 µg y digoxina 119,9 µg). La renovación adicional de la atmósfera interna en los SIT, como otro factor evaluado, indujo un mayor estrés oxidativo asociado a un aumento de la síntesis de digitoxina. Estos resultados permitieron realizar el escalado en SIT de 5L de capacidad y diseñar un esquema de producción como estrategia para la obtención de cardenólidos in vitro. Por otra parte, se demostró que la elicitación es una estrategia posible para el incremento de cardenólidos en brotes de D. purpurea. El SilioPlant® (0,01 g.L-1) incrementó la síntesis de cardenólidos en los brotes multiplicados in vitro en 3,6 y 6,9 veces el contenido de digoxina y digitoxina respectivamente, en comparación con el control. Se desarrolló un protocolo para la transformación genética con la cepa de A. tumefaciens C58C1RifR (pMP90) con el vector de transformación pTJK136. La presencia de los transgenes fue detectada mediante PCR y la hibridación por Southern en las líneas regeneradas. El número de insertos del nptII osciló entre una y dos. En el futuro, se requerirá de estudios relacionados con los mecanismos de síntesis de cardenólidos e identificación de enzimas específicas que permitan un incremento en la eficiencia de las alternativas estudiadas. ÍNDICE 1. INTRODUCCIÓN……………………………………………………………………………. 1 2. REVISIÓN BIBLIOGRÁFICA………………………………………………………………. 6 2.1 Digitalis purpurea L………………………………..………………………………………. 6 2.1.1 Origen, distribución y clasificación taxonómica…………………………..…………… 6 2.1.2 Descripción botánica……………………………….…………………………………… 6 2.1.3 Importancia farmacológica…………………………………………………………..…. 7 2.2 Cardenólidos .…………………………………………………………………...……..…….. 7 2.2.1 Estructura de los cardenólidos ………………………………………………..…….…… 7 2.2.2 Biosíntesis de cardenólidos………………………………………………………………. 8 2.2.3 Cultivo in vitro de Digitalis spp. y producción de cardenólidos………………………… 10 2.3 Producción in vitro de metabolitos secundarios………………………………………………. 11 2.3.1 Cultivo in vitro para la producción de metabolitos secundarios…………………………. 11 2.3.2 Escalado de la producción………………………………………………………………… 13 2.3.3 Sistemas de inmersión temporal……….……….…………………………………………. 15 2.3.4. Empleo de elicitores para la producción de metabolitos secundarios....…………………. 19 2.4 Transformación genética………………..…………………………..…………………………. 20 2.4.1 Ingeniería metabólica………..…….……………………………..……………………….. 20 2.4.2 Transformación genética mediada por Agrobacterium tumefaciens….…………...………. 21 2.4.3 Transformación genética en Digitalis spp. …………………..…………………………… 24 3. MATERIALES Y MÉTODOS………………………………………………………..…………. 26 3.1. Producción de biomasa de Digitalis purpurea en sistemas de inmersión temporal ..………... 30 3.1.1 Efecto de la densidad de inóculo……………….…………………………………………. 32 3.1.2 Efecto de la frecuencia de inmersión……………….……………………..……………… 32 3.1.3 Efecto de la renovación de la atmósfera…………………………….…………………….. 32 3.1.4 Escalado de la producción de biomasa ………………….......……………………………. 33 3.2 Efecto de elicitores en el cultivo de brotes de D.
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