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Evolución Termal Del Polimorfismo

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Evolución Termal Del Polimorfismo Departamento de Genética y de Microbiología Facultad de Ciencias EVOLUCIÓN TERMAL DEL POLIMORFISMO CROMOSÓMICO Y LA MORFOMETRÍA DEL ALA DE UNA POBLACIÓN EXPERIMENTAL DE Drosophila subobscura Walkiria Janneth Céspedes Vigoya 2006 Memoria presentada por la licenciada en Biología Walkiria Janneth Céspedes Vigoya para la obtención del titulo de Doctora en Ciencias Esta tesis ha sido realizada gracias a una Beca para Formación de Personal Investigador (FPI) del Ministerio de Ciencia y Tecnología, concedida por el periodo de Octubre 2001 a septiembre del 2005. Los Doctores Mauro Santos y Antonio Fontdevila, Catedráticos del Departamento de Genética y Microbiología de la Facultad de Ciencias de la Universidad Autónoma de Barcelona, Certifican que: Walkiria Janneth Céspedes Vigoya ha llevado a cabo bajo su dirección el trabajo de investigación realizado en el Departamento de Genética y de Microbiología de la Facultad de Ciencias de la Universidad Autónoma de Barcelona que ha llevado a la elaboración de la tesis Doctoral titulada: Evolución Termal del Polimorfismo Cromosómico y la Morfometría del ala de una población experimental de Drosophila subobscura Dr. Mauro Santos Dr. Antonio Fontdevila Agradecimientos Agradezco a los Doctores Mauro Santos y Antonio Fontdevila por integrarme a su equipo de trabajo durante estos 4 años, por el asesoramiento, dirección del trabajo y por su contribución en el enriquecimiento de mi aprendizaje de la biologia evolutiva. Al Doctor Joan Balanyà por su participación y orientación en la lectura de los polimorfismos cromosómicos. A Todo el Grupo de Biologia Evolutiva por su calida acogida, por el tiempo dedicado en nuestras largas conversaciones por aclarar aspectos y conceptos evolutivos, por sus criticas constructivas y sus aportaciones en cada uno de los distintos momentos del desarrollo del trabajo, y por su buen sentido del humor que hizo divertido algunos momentos difíciles del trabajo y de mi vida personal. Por ultimo, agradezco al Ministerio de Ciencia y Tecnología por la concesión de una beca FPI (FP2000-7001). A la Direcció General de Recerca (Generalitat de Catalunya) y la Fundación Ramón Areces, que financiaron el proyecto. A los meus Amics Per tot el que me’heu donat EVOLUCIÓN TERMAL DEL POLIMORFISMO CROMOSÓMICO Y LA MORFOMETRÍA DEL ALA DE UNA POBLACIÓN EXPERIMENTAL DE Drosophila subobscura Pagina 1. INTRODUCCION…………………………………………………………….. 1 1.1. La especie Drosophila subobscura……………………………………........... 1 1.1.1. Distribución geográfica………………………………………………….......... 4 1.1.2. Los cromosomas y sus inversiones…………………………………………… 5 1.1.3. Polimorfismos cromosómicos de D. subobscura………………………........... 9 1.2. La colonización de América por D. subobscura……………………………. 11 1.3. Carácter adaptativo de las inversiones cromosómicas……………………. 16 1.4. Variación del polimorfismos cromosómicos en D. Subobscura…………… 17 1.4.1. Clinas latitudinales de tamaño corporal en D. Subobscura…………………… 21 1.4.2. Relación entre temperatura y características morfométricas………………….. 22 1.4.3. Relación entre polimorfismo cromosómico y tamaño del ala………………… 23 1.5. Variación fenotípica…………………………………………………………. 24 1.5.1. Control de la variación fenotípica…………………………………………….. 25 1.5.1.1. Canalización……………………………………………………………........... 25 1.5.1.2. Estabilidad del desarrollo………………………………………………....... 27 1.5.1.3. Plasticidad………………………………………………………………….. 28 viii 1.5.2. Análisis de asimetría…………………………………………………………... 29 2. OBJETIVOS………………………………………………………………….. 32 2.1. Objetivo General……………………………………………………………... 32 2.2. Objetivos Específicos………………………………………………………… 32 3. ARTICULOS…………………………………………………………………. 34 3.1. Artículo 1………………………………………………………………........... 35 3.2. Artículo 2………………………………………………………………........... 54 3.3. Artículo 3………………………………………………………………........... 69 3.4 Artículo 4………………………………………………………………........... 89 4. DISCUSIÓN…………………………………………………………………... 117 4.1. Evolución termal de tamaño y forma del ala……………………………….. 117 4.1.1. Diversidad cromosómica………………………………………………………. 117 4.1.2. Variación en las frecuencias de los ordenamientos cromosómicos……............ 118 4.1.3 Evolución termal de las características del ala…………………………............ 119 4.1.3.1. Tamaño del ala…………………………………………………………............ 119 4.1.3.1.1. Tamaño celular y el índice del número celular…………………………........... 120 ix 4.1.3.1.2. Relación entre el polimorfismo de inversión y la variación clinal del tamaño del ala………………………………………………………………………….. 120 4.1.3.2. Forma del ala……………………………………………………………........... 121 4.1.3.2.1. Tasas de divergencia genética para forma del ala…………………………....... 122 4.1.3.2.2. Relación entre el polimorfismo de inversión y el índice de forma del ala…….. 123 4.2. Análisis de los efectos de la variación genética en las dimensiones del ala, en líneas isocromosómicas del cromosoma O………………......................... 124 4.2.1. Variación y asimetría en tamaño del ala………………………………………. 124 4.2.2. Variación y asimetría en la forma del ala………………………………............ 125 4.2.2.1. Patrones de la variación de la forma del ala………………………………........ 126 5. CONCLUSIONES……………………………………………………………. 128 REFERENCIAS……………………………………………………………… 129 x INDICE DE TABLAS Y FIGURAS Tabla 1: División en subgrupos del grupo obscura…………………………….......... 3 Tabla 2: Ordenaciones cromosómicas en América……………………………........... 14 Figura 1: Vista dorsal y lateral de Drosophila subobscura…………………………… 2 Figura 2: Distribución geográfica de Drosophila subobscura………………………... 4 Figura 3: Preparación de los cromosomas politenicos de Drosophila subobscura (5 acrocentricos y 1 puntiforme)………………………………………………. 5 Figura 4: Mapa de los cromosomas de Drosophila subobscura, realizada por Kunze- Muhl y Muller, y actualizada por Krimbas (1993) con nuevas 7 inversiones…………………………………………………………….......... Figura 5: Inversión A2.................................................................................................... 8 Figura 6: Ordenamiento O7…………………………………………………………… 9 Figura 7: Ordenamiento O3+4+7....................................................................................... 9 Figura 8: Ordenamiento E1+2+9…………………………………………………........... 9 Figura 9: Ordenamiento O3+4…………………………………………………………. 9 Figura 10: Ordenamiento Ast…………………………………………………………… 15 Figura 11: Ordenamiento Jst……………………………………………………………. 15 Figura 12: Ordenamiento J1…………………………………………………………….. 15 Figura 13: Ordenamiento Ust…………………………………………………………… 15 xi Figura 14: Ordenamiento U1+2…………………………………………………………. 15 Figura 15: Ordenamiento U1+2+8…………………………………………………........... 15 Figura 16: Ordenamiento Est…………………………………………………………… 15 Figura 17: Ordenamiento E1+2+9+3……………………………………………………… 15 Figura 18: Ordenamiento E1+2+9+12………………………………………………........... 16 Figura 19: Ordenamiento Ost…………………………………………………………… 16 Figura 20: Ordenamiento O3+4+2…………………………………………………........... 16 Figura 21: Ordenamiento O3+4+8…………………………………………………........... 16 Figura 22: Ordenamiento O5…………………………………………………………… 16 Figura 23: Canalización por norma de reacción............................................................... 26 Figura 24: Canalización por variación aproximada......................................................... 26 Figura 25: Distribuciones mas comunes de R-L en organismos de simetría bilateral: a) asimetría fluctuante (FA) b) Asimetría direccional (DA) y c) antisimetría 29 (Platykurtic o bimodal)………………………………................................... xii ABREVIATURAS Ba: Gen dominante Bare que determina el carácter del número de macroquetas. Número que normalmente está reducido en el mutante. Sin embargo las fosas de las cerdas están siempre presentes. Es letal en condición de homocigosis. ch: Gen para el carácter de ojos color cherry CS: Centroid Size – Tamaño del centroide cu: Gen para el carácter de alas curvas (curled) DA: Directional Asymmetry - Asimetría direccional DI: Developmental instability – Inestabilidad del desarrollo E: Eastern FA: Fluctuating Asymmetry - Asimetría fluctuante L1: Longitud del segmento basal o proximal de la vena longitudinal IV del ala L2: Longitud del segmento distal de la vena longitudinal IV del ala L-R: Left-Right – Izquierda - Derecha N: North PCs: Componentes principales S: South st: Ordenamiento cromosómico estándar Va: Gen Varicose, gen dominante que determina el carácter carácter del engrosamiento irregular en la unión de las venas alares y la bifurcación en las venas de tamaño irregular corto. Es letal en condición de homocigosis. W: West WL: Wing Length– Longitud total del ala WS: Wing Shape – forma del ala xiii 1. INTRODUCCION Diversas especies de Drosophila son colonizadoras, lo cual ha permitido utilizarlas como especies modélicas en el estudio de los importantes procesos evolutivos que acompañan la colonización. En el estudio de la colonización de América por D. subobscura se ha demostrado que la expansión ha sido muy rápida y que ha conducido al establecimiento del mismo tipo de clinas latitudinales para caracteres cromosómicos y morfológicos presentes en el Viejo Mundo (poblaciones paleárticas), lo que demuestra la plasticidad evolutiva de la especie a este nivel de variabilidad genética y la capacidad de su adaptación rápida según un gradiente latitudinal. Las clinas latitudinales para ordenaciones cromosómicas aparecieron en tan sólo 3 años en Suramérica y 4 años en Norteamérica después del inicio de la colonización, aproximadamente hace 25 años. Las áreas colonizadas tienen una situación geográfica equivalente en ambos hemisferios, y las condiciones climáticas son paralelas a las del área de distribución en la región paleártica (Prevosti
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