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Plazas Ávila UNIVERSITAT POLITÈCNICA DE VALÈNCIA Departament de Biotecnologia Tesis para la obtención del Título de Doctora en Biotecnología CARACTERIZACIÓN Y MEJORA GENÉTICA DE LA BERENJENA (S. melongena L.) PARA COMPUESTOS BIOACTIVOS AUTORA: MARÍA DE LA O PLAZAS ÁVILA DIRIGIDA POR: JAIME PROHENS TOMÁS SANTIAGO VILANOVA NAVARRO ISABEL ANDÚJAR PÉREZ Valencia, 2015 “ Lo que no se empieza, nunca tendrá un final” Johan Wolfgang von Goethe A Mario y a Vega A mis padres A Jorge Agradecimientos Tengo tantas gracias que dar que no se por donde empezar. Así que, para ser justa debería empezar dando las gracias al Universo, por aquel día hace casi 15 años que comiendo en agrícolas me encontré un cartel donde ponía: “se busca estudiante para trabajar con Pepino Dulce, preguntar por Jaime Prohens”. Esa fue la primera vez que escuché su nombre, y nada presagiaba que iba a ser parte de mi vida… espero que para siempre! Jaime, jamás hubiera podido hacer esto sin ti, además, jamás me lo habría planteado. Gracias por estar siempre ahí, dispuesto a ayudarme y a aconsejarme siempre que lo he necesitado. Has formado parte de muchas cosas importantes que han pasado en mi vida, tanto profesional como personalmente. Tengo tanto que agradecerte que podría dedicarte esto solo a ti, pero no seria justo ;) Santi que voy a decirte, siempre con una sonrisa y animándome cada una de las veces que te he necesitado. Me encanta saber que tu buen humor y tu supercerebro van a estar ahí, siempre que lo necesite. Isa, la más reciente incorporación al grupo y no por ello menos importante, gracias por ser mi complemento, por ayudarme en todo y por estar siempre disponible. Pietro, mi italiano preferido, con el que siempre puedo hablar de política o de cualquier otra cosa, gracias por estar ahí. Javi, bienvenido a los melongena, espero que todo lo que he pasado te ayude a llevarlo mejor cuando te toque a ti ;). María, aunque ahora estas un poco más lejos que sepas que te echo de menos. Dionís, mi “xiquet”, has traído la juventud y la inocencia de nuevo al laboratorio y, sobre todo, la motivación a dejarnos la carne ;) No menos importante sois toda la gente que ha pasado por el laboratorio berenjena con la que he compartido tantas cosas, tantas risas, tantos almuerzos y tanto buen rollo que ha hecho que siempre tuviera ganas de madrugar para llegar al trabajo. Han pasado tantas personas por aquí, en todo el tiempo que llevo en el COMAV, que seguro si intentara nombrar a todas, me olvidaría de alguien, así que daros por recordadas. De cada una he aprendido algo y de todas me llevo algún recuerdo. Esto ha sido como la ONU… y más que va a ser! Muchas gracias a todo el grupo del COMAV con el que he compartido muy buenos ratos y sobre todo, a quienes habéis estado ahí cuando en algún momento he necesitado desahogarme: Ana Fita, Adrián, Nuri, Álvaro, Inma, Ali, las Mª José, Tere, Gloria, Carlos, Paco, Pascual, Vero, Estela, Loles… Qué les podría decir a mis chicas -mi trío-. Habéis estado ahí animándome durante todo el proceso, dándome sabios consejos para realizar esta Tesis y recordándome que SOY LA MEJOR, jajaja. Gracias chicas, en cuanto termine nos vamos juntas al fin del mundo. Durante este tiempo ha habido muchos momentos buenos, algunos regulares y otros malos; gente a la que me hubiera gustado tener aquí ahora y que me estará viendo desde algún lugar, dándome su apoyo, a la que echo mucho de menos y que me da fuerzas para seguir. Con esta Tesis le he quitado tiempo a mi familia, a Vega, a Mario y a Jorge. Habéis tenido que mantener mis nervios y también pasar muchos momentos sin mí, que prometo recuperar. Sois mi fuerza, mi apoyo, mi ilusión, mis amores, lo sois todo y os quiero más que a nada. Gracias Pedro, gracias Virtudes siempre habéis sido mi apoyo y mi máxima motivación a superarme. Gracias César y gracias Laura por estar siempre a mi lado. A Jaime y Mª Carmen, que tanto me habéis ayudado en esta etapa de “Estudios y Política”, sin esta gran familia que somos, habría sido imposible. También quiero dar las gracias a mis amigos y amigas por ser mis incondicionales, a Marcos y a Isa por la portada y a María Eugenia por la frase, y como no, a “la plaza” por las buenas conversaciones y el desahogo, que tantas veces me han alimentado y animado a seguir ;) Gracias a todos y a todas, OS QUIERO! Índice Índice ÍNDICE SUMMARY 1 RESUMEN 5 RESUM 9 1. INTRODUCCIÓN 13 1.1. Breeding vegetables with improved bioactive properties 15 1.2. La berenjena como cultivo objetivo para la mejora de compuestos bioactivos 33 1.3. Diversidad en berenjena como materia prima para la mejora 47 1.4. Breeding for chlorogenic acid content in eggplant: interest and prospects 67 2. OBJETIVOS 91 3. RESULTADOS 95 3.1. Diversidad en berenjena común para compuestos bioactivos y caracteres relacionados 97 3.1.1. Diversity and relationships in key traits for functional and apparent quality in a collection of eggplant: fruit phenolics content, antioxidant activity, polyphenol oxidase activity, and browning 99 3.2. Diversidad en berenjenas escarlata y gboma para una mejora integral: caracteres morfoagronómicos y compuestos bioactivos 131 3.2.1. Conventional and phenomics characterization provides insight into the diversity and relationships of hypervariable scarlet (Solanum aethiopicum L.) and gboma (S. macrocarpon L.) complexes 133 3.2.2. Reducing capacity, chlorogenic acid content and biological activity in a collection of scarlet (Solanum aethiopicum) and gboma (S. macrocarpon) eggplants 167 3.3. Hibridación interespecífica para la mejora del contenido en compuestos bioactivos de la berenjena 199 3.3.1. Characterization of interspecific hybrids and first backcross generations from crosses between two cultivated eggplants (Solanum melongena and S. aethiopicum Kumba group) and implications for eggplant breeding 201 3.3.2. Genetic diversity in morphological characters and phenomic acids content resulting from an interspecific cross between eggplant, Solanum melongena, and its wild ancestor (S. incanum) 243 4. DISCUSIÓN GENERAL 279 5. CONCLUSIONES 305 Summary SUMMARY Fruits and vegetables contain bioactive compounds beneficial for human health. The development of varieties with a higher content in these compounds is of interest, as it contributes to satisfying an increasing demand by consumers of products with functional activity. Among other vegetables, eggplant (Solanum melongena) has a high antioxidant activity, mostly derived from its high content in polyphenols, and it has been demonstrated to have beneficial effects for human health. Amongst the phenolic compounds of eggplant, chlorogenic acid outstands, as it is the most abundant phenolic compound in this crop and presents multiple beneficial properties for health. This Doctoral Thesis deals with the characterization and breeding of eggplant in order to obtain relevant information and plant material for the development of eggplant varieties with a higher content in bioactive compounds, in particular, polyphenols, making use of the intraspecific and interspecific variation. On the other hand, an integral breeding approach must take into account not only the trait to be improved, but also those traits of interest for the success of a variety and, in consequence, we have studied other traits related to the increase in the phenolic content, like fruit browning, and also other traits of general interest for breeding. In the first part of this Doctoral Thesis we focus on the study of the diversity of common eggplant and related species in traits of agronomic interest. The objective is to evaluate the diversity, identify sources of variation and to study relationships among traits. In a first study, we evaluate a collection of traditional eggplant varieties, in which we have found a high diversity for functional quality traits and browning. In this study we found that the content in chlorogenic acid is positively correlated with the antioxidant activity and the correlation with browning is low, demonstrating that it is feasible to select eggplant varieties with high content in chlorogenic acid and moderate browning. We also found that, even with a low polyphenol oxidase activity, there may be a significant browning, suggesting that polyphenol oxidase activity (PPO) is not the limiting factor for browning in the studied collection. With the aim of increasing the genetic diversity for breeding eggplant for bioactive compounds and other traits of importance, we have studied the 1 diversity in a collection of scarlet (S. aethiopicum) and gboma (S. macrocarpon) eggplants. The morphological characterization with conventional descriptors and phenomic tools (Tomato Analyzer) has allowed us to study the relationships among the different cultivar groups and wild relatives and to determine that scarlet and gboma eggplant complexes are hypervariable. In this collection we have also studied the reducing capacity and the content in chlorogenic acid, and we have found a huge variability. In general, scarlet eggplant presents relatively low contents, while gboma eggplant, in particular its wild ancestor S. dasyphyllum, presents very high values. In macrophage cell cultures we have also found that the varieties with higher content in chlorogenic acid also display a greater inhibition of nitric oxide (NO) production indicating beneficial properties for health. In the second part of this Doctoral Thesis we have evaluated the interest of interspecific hybridization for eggplant breeding, in particular for the content in bioactive compounds. We have obtained two families, including backcrosses, between common eggplant (S. melongena) on one side and cultivated scarlet eggplant (S. aethiopicum) and the wild relative S. incanum on the other. The results show that fertility of materials derived from the hybridization between S. melongena and S. aethiopicum is low and that there is a low efficiency in the backcrosses to S.
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