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Departament De Biologia Funcional I Antropologia Física DEPARTAMENT DE BIOLOGIA FUNCIONAL I ANTROPOLOGIA FÍSICA EVALUACIÓN DE MECANISMOS DE RESISTENCIA A INSECTICIDAS EN FRANKLINIELLA OCCIDENTALIS (PERGANDE): IMPLICACIÓN DE CARBOXILESTERASAS Y ACETILCOLINESTERASAS NEUS LÓPEZ SOLER UNIVERSITAT DE VALÈNCIA Servei de Publicacions 2008 Aquesta Tesi Doctoral va ser presentada a València el dia 26 de juny de 2008 davant un tribunal format per: - D. Rafael Martínez Pardo - D. Félix Ortego Alonso - D. Juan Ferré Manzanero - D. Francisco José Beitia Crespo - D. Juan Javier Díaz Mayans Va ser dirigida per: Dª. Mª Dolores Garcerá Zamorano Dª. Amelia Cervera Olagüe ©Copyright: Servei de Publicacions Neus López Soler Depòsit legal: I.S.B.N.: 978-84-370-7227-2 Edita: Universitat de València Servei de Publicacions C/ Artes Gráficas, 13 bajo 46010 València Spain Telèfon: 963864115 Facultat de Ciències Biològiques Departament de Biologia Funcional i Antropologia Física Evaluación de mecanismos de resistencia a insecticidas en Frankliniella occidentalis (Pergande): implicación de carboxilesterasas y acetilcolinesterasas Neus López Soler Burjassot, València 2008 i Tesis presentada por Neus López Soler para optar al grado de Doctor en Ciencias Biológicas por la Universitat de València. Fdo. Neus López Soler Tesis dirigida por: Mª Dolores Garcerá Zamorano, Doctora en Ciencias Biológicas y Catedrática de Fisiología de la Universitat de València. Amelia Cervera Olagüe, Doctora en Ciencias Biológicas. Fdo. Mª Dolores Garcerá Zamorano Fdo. Amelia Cervera Olagüe La investigación presentada en esta Tesis se realizó en la Unidad de Fisiología Animal del Departament de Biologia Funcional i Antropologia Física de la Facultat de Ciències Biològiques de la Universitat de València, gracias al disfrute de una Beca del Programa Nacional de Formación de Profesorado Universitario (FPU) del Ministerio de Educación y Ciencia. ii A Mari i Santi iii I després de creure tant t´ arribaren tots els dubtes, tenia raó el filòsof: pressuposa la certesa el mateix fet de dubtar (I després de creure tant, RAIMON) iv Agradecimientos Encara no sé ben cert perquè vaig decidir fer el doctorat, però supose que en part per inquietud i en part per atzar. En aquell moment tan sols em plantejava si m’ agradaria treballar en un laboratori, però finalment he acabat esbrinant moltísimes més coses d´ aquest món que anomenem Ciència, i també d´ altres. Primerament vaig descobrir el que volia dir un dia de treball “frenètic” al laboratori, per descomptat, de la mà de les meues companyes (quasi sempre han estat companyes). A poc a poc vaig ser capaç de suportar-los, assumir-los i inclús organitzar-los i gaudir-los. Després vaig començar a saber per a que servia això de “fitxar la bibliografia”, i per descomptat la seua cerca, lectura, enteniment, etc. Això em va conduir a donar un pas més enllà i tractar de dissenyar experiments, millorar altres, començar amb alguna nova tècnica (llegint prèviament les “tedioses” instruccions dels aparells), anàlisi estadístic de dades, escriptura de treballs, i un llarg etcètera, incloent coses no tant directament relacionades (encara que en certs moments, inclús, més importants) com qüestions burocràtiques, d’informàtica, i un altre llarg etcètera. A banda he pogut gaudir del privilegi d´ haver treballat a altres centres d´ investigació (Rothamsted Research, Anglaterra, i Bayer CropScience, Alemanya), aprenent formes diferents de treball, organització, direcció, etc. De la mateixa manera, he pogut participar en projectes duts a terme en col·laboració amb altres grups, amb una major complexitat i diversitat. Paral·lelament, he pogut saber que vol dir això de tenir companys de treball, amb els quals aprens a compartir espais, treball, inquietuds, problemes, moments de descans... i fins i tot, una amistat. I també el procés a la inversa, compartint amb els meus amics i sobretot amb la meua família, tot allò relacionat amb el treball. És a dir, he començat a tenir nocions del v que vol dir barrejar les facetes laboral i personal, ambdues tant importants a les nostres vides. I encara que els que em coneixen saben que per a mi estar a l´ estranger no és una predilecció, també ha resultat ser una molt bona experiència personal per haver pogut conéixer gent “diferent”, situacions “estranyes”, etc. Tot en conjunt ha estat un llarg i intens camí, ple d´ esdeveniments, sentiments, i per descomptat, també, algun que altre entrebanc. Però el més important de tot heu estat totes les persones que m´ heu acompanyat en aquesta trajectòria, en totes les seues facetes i en tots els seus aspectes. Tot açò no haguera sigut possible sense la inestimable ajuda que he rebut. I com que aquest manuscrit és fonamentalment científic, vull agrair especialment a Ame i Loles la seua confiança i dedicació. Aquest treball és també de tots Vosaltres. vi Índice Índice 1. Introducción .......................................................................................1 1.1. El trips occidental de las flores, Frankliniella occidentalis ............6 1.1.1. Taxonomía y biogeografía .............................................6 1.1.2. Biología de Frankliniella occidentalis ..............................7 1.1.3. El trips como plaga: aspectos biológicos que inciden en las estrategias de control .........................................11 1.1.4. Incidencia de la plaga en España .................................13 1.2. Clasificación de los insecticidas ...............................................15 1.2.1. Inhibidores de acetilcolinesterasas (AChE) ....................16 1.2.2. Antagonistas de los canales de cloro dependientes del ácido -aminobutírico (GABA) .................................18 1.2.3. Moduladores de los canales de sodio ...........................18 1.3. Resistencia a insecticidas .......................................................19 1.3.1. Aspectos generales .....................................................19 1.3.2. Mecanismos de resistencia a insecticidas .....................23 1.3.2.1. Resistencia por comportamiento ......................24 1.3.2.2. Reducción de la penetración ...........................24 1.3.2.3. Incremento de la destoxificación metabólica.....25 1.3.2.4. Alteración de los puntos de acción de los insecticidas ....................................................30 1.3.3. Procedimientos experimentales usuales en el estudio de la resistencia ...........................................................34 1.4. Objetivos ................................................................................37 2. Estudio de isoenzimas de esterasas en poblaciones de campo de Frankliniella occidentalis y su relación con la resistencia ............39 2.1. Introducción ............................................................................41 2.2. Material y métodos ..................................................................44 vii 2.2.1. Insectos ......................................................................44 2.2.2. Preparación de las muestras ........................................45 2.2.3. Electroforesis en geles de poliacrilamida (PAGE) y tinción para actividad esterasa ......................................45 2.2.4. Optimización del ensayo α-naftil butirato .......................49 2.2.5. Actividad esterasa total ................................................50 2.2.6. Determinación de proteína ...........................................51 2.2.7. Análisis estadístico ......................................................51 2.3. Resultados .............................................................................53 2.3.1. Electroforesis en geles de poliacrilamida: tinción con α-naftil acetato .............................................................53 2.3.2. Optimización del ensayo α-naftil butirato .......................59 2.3.3. Actividad esterasa total: α-naftil acetato y α-naftil butirato .......................................................................61 2.3.4. Electroforesis en geles de poliacrilamida: tinción con α-naftil acetato y α-naftil butirato ...................................63 2.4. Discusión ................................................................................64 3. Análisis del efecto de diversos compuestos sinergistas sobre la actividad esterasa en Frankliniella occidentalis ..................71 3.1. Introducción ............................................................................73 3.2. Material y métodos ..................................................................76 3.2.1. Insectos ......................................................................76 3.2.2. Preparación de las muestras ........................................77 3.2.3. Compuestos químicos utilizados ...................................77 3.2.4. Inhibición de la actividad esterasa total por sinergistas ..78 3.2.5. Electroforesis en geles de poliacrilamida (PAGE) nativos e inhibición de esterasas por sinergistas ............79 3.2.6. Inhibición de la actividad esterasa mediante bioensayos de sinergistas ............................................80 3.2.7. Determinación de proteína ...........................................81 3.2.8. Análisis estadístico ......................................................81 viii Índice 3.3. Resultados .............................................................................83 3.3.1. Análisis de inhibición de la actividad esterasa total por sinergistas .............................................................83 3.3.2. Análisis de inhibición de isoenzimas de esterasas por sinergistas .............................................................86
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