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The Diversity of Agriculture and Human Culture Are Bound Together. in The “The diversity of agriculture and human culture are bound together. In the end it is up to all of us as governments and communities and individuals to prize diversity.” Cary Fowler and Pat Mooney ORIGEN, DISPERSIÓN Y DIVERSIDAD DEL CHIRIMOYO (ANNONA CHERIMOLA MILL.) EN EL CONTINENTE AMERICANO Memoria de tesis doctoral presentada por Nerea Larrañaga González para optar al grado de Doctora por la Universidad Politécnica de Madrid, programa de Doctorado en Biotecnología y Recursos Genéticos en Plantas y Microorganismos Asociados. Director: Dr. José Ignacio Hormaza Urroz, Profesor de Investigación. Septiembre de 2016 Universidad de Málaga, Consejo Superior de Investigaciones Científicas Instituto de Hortofruticultura Subtropical y Mediterránea La Mayora (IHSM-UMA-CSIC) 29750 Algarrobo –Costa (Málaga) José Ignacio Hormaza Urroz, Profesor de Investigación del Consejo Superior de Investigaciones Científicas en el Instituto de Hortofruticultura Subtropical y Mediterránea La Mayora y director del Departamento de Fruticultura, CERTIFICA que Nerea Larrañaga González, Licenciada en Biología y Máster en Recursos Fitogenéticos ha realizado en el Departamento de Fruticultura del Instituto de Hortofruticultura Subtropical y Mediterránea La Mayora (IHSM-UMA-CSIC), bajo su dirección, el trabajo de investigación que con el título “Origen, dispersión y diversidad del chirimoyo (Annona cherimola Mill.) en el continente americano” presenta para optar al grado de Doctora por la Universidad Politécnica de Madrid. Septiembre de 2016 Fdo. Dr. José Ignacio Hormaza Urroz Universidad Politécnica de Madrid Escuela Técnica Superior de Ingeniería Agronómica, Alimentaria y de Biosistemas Departamento de Biotecnología – Biología Vegetal 28040 Madrid Juan Bautista Martínez Laborde, profesor titular del Departamento de Biotecnología-Biología Vegetal de la Escuela Técnica Superior de Ingeniería Agronómica, Alimentaria y de Biosistemas de la Universidad Politécnica de Madrid AUTORIZA la presentación de la Tesis Doctoral titulada “Origen, dispersión y diversidad del chirimoyo (Annona cherimola Mill.) en el continente americano” de la doctoranda Nerea Larrañaga González realizada en el Departamento de Fruticultura del Instituto de Hortofruticultura Subtropical y Mediterránea La Mayora (IHSM-UMA-CSIC), para optar al grado de Doctora por la Universidad Politécnica de Madrid. Septiembre de 2016 Fdo. Dr. Juan Bautista Martínez Laborde Este trabajo se ha llevado a cabo en el Departamento de Fruticultura del Instituto de Hortofruticultura Subtropical y Mediterránea La Mayora, instituto mixto del Consejo Superior de Investigaciones Científicas y la Universidad de Málaga gracias a una ayuda predoctoral de Formación de Personal Investigador (FPI) (BES-2011-047451) del Ministerio de Economía y Competitividad de España, asociada al proyecto AGL2010-1514. Tribunal nombrado por el Magfco. y Excmo. Sr. Rector de la Universidad Politécnica de Madrid, el día Presidente: Vocal: Vocal: Vocal: Secretario: Suplente: Suplente: Realizado el acto de defensa y lectura de la tesis el día ……………de …………… de 2016 en la E.T.S. de Ingeniería Agronómica, Alimentaria y de Biosistemas de la Universidad Politécnica de Madrid. CALIFICACIÓN………. ……………….. PRESIDENTE ……………….. SECRETARIO ……………….. VOCALES Parte de los resultados de esta Tesis Doctoral se incluyen en los siguientes artículos publicados en revistas científicas recogidas en el SCI y capítulo de libro: Larranaga N, Hormaza JI (2015) DNA barcoding of perennial fruit tree species of agronomic interest in the genus Annona (Annonaceae). Frontiers in Plant Science, 6: 589 (Anexo 1). Larranaga N, JI Hormaza (2016) Advances in genetic diversity analysis in fruit tree crops. Lüttge U, Cánovas FM, Matyssek R. Progress in Botany 77. Springer International Publishing, Switzerland, 245-264 (Anexo 2). Larranaga N, Albertazzi F, Fontecha G, Palmieri M, Rainer H, van Zonneveld M, Hormaza, JI (2016) A Mesoamerican origin of cherimoya (Annona cherimola Mill.), implications for the conservation of plant genetic resources. Molecular Ecology (en revisión). Y presentados en los congresos Larrañaga N, Albertazzi F, Fontecha GA, Palmieri M, van Zonneveld M, Hormaza JI (2013) Cherimoya (Annona cherimola Mill.) genetic diversity and structure across America: possible Central American origin. V International congress on Annonaceae. Botucatu, Brasil, 19-23 de agosto. Larrañaga N, van Zonneveld M, Albertazzi F, Fontecha GA, Palmieri M, Hormaza JI (2014) New insights on the center of origin of cherimoya (Annona cherimola Mill., Annonaceae) combining molecular tools and Geographic Information Systems. XXII Plant & Animal Genome conference. San Diego, EEUU, 11-15 de enero. AGRADECIMIENTOS Son muchas las personas que de alguna manera han contribuido a la concreción de este trabajo de investigación y a las que quiero dar las gracias. En primer lugar a agradezco a Iñaki Hormaza, director de esta tesis, por haberme dado la oportunidad, la confianza y gran parte de su tiempo; por lo mucho que he aprendido tanto en lo científico como en lo personal. A todo el grupo del laboratorio de Fruticultura que desde el primer día me hicieron sentir parte de él; gracias a Yolanda, Librada, Carolina, Jorge L., Alicia, The, Celeste, Inma, Marta y Rosa. Y por supuesto a Vero, compañera y amiga. Gracias a Juan B. Martínez, tutor académico de esta tesis, siempre dispuesto a recibirme y echarme un cable con los trámites administrativos. Gracias a Federico Albertazzi, Gustavo Fontecha, Maarten van Zonneveld y Lars Chatrou por acogerme en sus respectivas instituciones (y ciudades) durante las estancias internacionales en San José, Tegucigalpa, Cali y Wageningen. Han sido experiencias enriquecedoras en todos los sentidos. No me olvido de Mariela, Clara, Elisa, Marlon, María, Óscar, Melisa, Dani, Jenifer y el resto del equipo que me recibió durante esos meses. También agradezco las colaboraciones con Margarita Palmeri (Guatemala) y Jorge Andrés (México). Agradezco al resto de compañerxs del Centro y del cuartito, con quienes hemos compartido mucho más que un lugar de trabajo: María, Anabel, Olaya, Yazmín, Jorge, Kike, Cañi, Mariola, Willy, Reme, Rida, Sara, Jota, Paco, Elisa, Eduardo, Pedro, Martin, Anelise, Davinia, Paola, Rima, Isa, Elvira, Antonio, Luis, Irene y tantos otrxs. Y en general, a todo el personal de la Mayora, con especial mención al departamento de Informática por su ayuda en lo referente a programas y otros dramas tecnológicos, a los compañerxs de campo y de administración. Gracias también a Antonio Matas por darme acceso al servidor de R de la UMA, a Silvia Martín de la UPM por la orientación con toda la documentación y a Ricardo Ontiveros por su asesoría con R. Gracias a mis amigxs de siempre, porque nunca han dejado de acompañarme y de tratar de entender por qué el estudio de la diversidad genética del chirimoyo es tan importante. A mis padres: Sabin (quien ha hecho la portada) y Ángeles (quien ha dibujado la chirimoya), y a mi hermana Leire, que me apoyan incondicionalmente. A mi pareja, David, quien conoce cada punto y coma de este trabajo y está a mi lado con cada decisión. A Trufa, mi perrita, que se ha pasado las tardes de los últimos meses sin apenas salir y tumbada junto a mi mesa. Finalmente, quiero dar las gracias a las instituciones responsables de esta Tesis: al CSIC, a la Universidad Politécnica de Madrid y a la beca del Ministerio de Economía y Competitividad que me fue otorgada. También a todas las entidades públicas y grupos de investigación donde he recibido la formación académica y científica previa. RESUMEN En este trabajo se ha profundizado en las razones que pueden explicar la distribución actual de la diversidad genética del chirimoyo (Annona cherimola Mill.), una especie frutal adaptada a climas subtropicales, con claras implicaciones para su uso y conservación. Gracias a un estudio de código de barras, se pudo comprobar el poder de discriminación del gen cloroplastídico maturasa K (matK) entre las especies con mayor importancia agronómica del género. Además, sobre los polimorfismos de nucleótido único (SNPs) interespecíficos, se diseñaron marcadores que permitieron la identificación mediante presencia o ausencia de bandas en geles de electroforesis de dichas especies tras PCR. El marcador específico del chirimoyo se utilizó para amplificar un gran número de muestras y estudiar la variabilidad intraespecífica respecto a los SNPs elegidos. Se verificó la existencia de dos haplotipos con diferenciación espacial: (1) con presencia únicamente en Centroamérica y (2) en Centroamérica y resto de orígenes de las accesiones del banco de germoplasma del IHSM- UMA-CSIC. A través de un estudio filogenético, basado en cinco loci cloroplastídicos, se determinó que esta especie conforma un grupo monofilético con otras especies cercanas, las cuales tienen una distribución exclusivamente centroamericana y caribeña. De entre las accesiones de chirimoyo incluidas en este ensayo, la mexicana y hondureñas (que presentan el haplotipo único de Centroamérica para matk) se posicionaron en la base del clado que reúne a todas las accesiones de la especie y no se distinguieron de las muestras de Annona pruinosa. La aproximación espacial para analizar diferentes índices de diversidad genética a partir de una base de datos final de 1843 accesiones de chirimoyo y nueve marcadores microsatélite, mostró como los valores en la zona centroamericana (preferentemente Honduras y Guatemala) fueron más altos en comparación con el norte de Perú, donde se observaron los valores más altos de
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    LISTE INTERNATIONALE DES NOMS PROTÉGÉS (également disponible sur notre Site Internet : www.IFHAonline.org) INTERNATIONAL LIST OF PROTECTED NAMES (also available on our Web site : www.IFHAonline.org) Fédération Internationale des Autorités Hippiques de Courses au Galop International Federation of Horseracing Authorities 15/04/13 46 place Abel Gance, 92100 Boulogne, France Tel : + 33 1 49 10 20 15 ; Fax : + 33 1 47 61 93 32 E-mail : [email protected] Internet : www.IFHAonline.org La liste des Noms Protégés comprend les noms : The list of Protected Names includes the names of : F Avant 1996, des chevaux qui ont une renommée F Prior 1996, the horses who are internationally internationale, soit comme principaux renowned, either as main stallions and reproducteurs ou comme champions en courses broodmares or as champions in racing (flat or (en plat et en obstacles), jump) F de 1996 à 2004, des gagnants des neuf grandes F from 1996 to 2004, the winners of the nine épreuves internationales suivantes : following international races : Gran Premio Carlos Pellegrini, Grande Premio Brazil (Amérique du Sud/South America) Japan Cup, Melbourne Cup (Asie/Asia) Prix de l’Arc de Triomphe, King George VI and Queen Elizabeth Stakes, Queen Elizabeth II Stakes (Europe/Europa) Breeders’ Cup Classic, Breeders’ Cup Turf (Amérique du Nord/North America) F à partir de 2005, des gagnants des onze grandes F since 2005, the winners of the eleven famous épreuves internationales suivantes : following international races : Gran Premio Carlos Pellegrini, Grande Premio Brazil (Amérique du Sud/South America) Cox Plate (2005), Melbourne Cup (à partir de 2006 / from 2006 onwards), Dubai World Cup, Hong Kong Cup, Japan Cup (Asie/Asia) Prix de l’Arc de Triomphe, King George VI and Queen Elizabeth Stakes, Irish Champion (Europe/Europa) Breeders’ Cup Classic, Breeders’ Cup Turf (Amérique du Nord/North America) F des principaux reproducteurs, inscrits à la F the main stallions and broodmares, registered demande du Comité International des Stud on request of the International Stud Book Books.
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