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Naturalis Repositorio Institucional Universidad Nacional De La Plata Facultad De Ciencias Naturales Y Museo Naturalis Repositorio Institucional Universidad Nacional de La Plata http://naturalis.fcnym.unlp.edu.ar Facultad de Ciencias Naturales y Museo Citogenética básica y evolutiva de Heterópteros fitófagos de interés agroeconómico de la Argentina Bressa, María José Doctor en Ciencias Naturales Dirección: Larramendy, Marcelo Luis Co-dirección: Papeschi, Alba Graciela Facultad de Ciencias Naturales y Museo 2003 Acceso en: http://naturalis.fcnym.unlp.edu.ar/id/20120126000135 Esta obra está bajo una Licencia Creative Commons Atribución-NoComercial-CompartirIgual 4.0 Internacional Powered by TCPDF (www.tcpdf.org) UNIVERSIDAD NACIONAL DE LA PLATA FACULTAD DE CIENCIAS NATURALES Y MUSEO Cátedra de Citología Tesis presentada para obtener el Grado de Doctor en Ciencias Naturales CITOGENETICA BASICA Y EVOLUTIVA DE HETEROPTEROS FITOFAGOS DE INTERES AGROECONOMICO DE LA ARGENTINA María José Bressa Codirectora: Director: Dra. Alba Graciela Papeschi Dr. Marcelo L. Larramendy La Plata, 2003 A Alejandra, Citogenética Básica y Evolutiva de Heterópteros Fitófagos de Interés Agroeconómico de la Argentina I TABLA DE CONTENIDOS RESUMEN IV-VI LISTA ORIGINAL DE PUBLICACIONES VII INTRODUCCION 1 1. Generalidades de Heteroptera 1 1.1. Clasificación y filogenia 1-2 1.2. Características biológicas 2-5 1.3. Importancia agroeconómica de especies de las familias Coreidae, Largidae, Lygaeidae, Pyrrhocoridae y Rhopalidae para la Argentina 5-7 2. Antecedentes Citogenéticos de Heteroptera 7-8 2.1. Cromosomas holocinéticos 8-11 2.2. Comportamiento mitótico de los cromosomas holocinéticos 11 2.3. Orientación de los cromosomas meióticos 11-14 2.4. Comportamiento meiótico de los autosomas 14-17 2.5. Sistemas cromosómicos de determinación del sexo 17-18 2.6. Comportamiento meiótico de los cromosomas sexuales 18-19 2.7. Cromosomas m 19-21 2.8. Cromosomas supernumerarios 21-22 2.9. Comportamiento meiótico de univalentes autosómicos 22-23 2.10. Mecanismos de evolución del cariotipo en Heteroptera 23-25 3. Antecedentes Moleculares y Citogenético-Moleculares de Heteroptera 26-28 4. Generalidades de las Familias Coreidae, Largidae, Lygaeidae, Pyrrhocoridae y Rhopalidae 28 4.1. Características biológicas 28-30 4.2. Antecedentes citogenéticos 30-31 OBJETIVOS DEL ESTUDIO 32 MATERIALES Y METODOS 33 5. Colección del Material 33-36 6. Citogenética Básica 37 6.1. Fijación 37 6.2. Determinación taxonómica 37-38 6.3. Disección de las gonadas 38 Citogenética Básica y Evolutiva de Heterópteros Fitófagos de Interés Agroeconómico de la Argentina II 6.4. Confección de preparados citogenéticos con hematoxilina férrica acética 38 6.5. Confección de preparados citogenéticos para posterior tinción con 4'6-diamidino-2-fenilindol (DAPI), Feulgen e impregnación argéntica 39 6.6. Análisis citogenético 39-40 6.7. Análisis estadístico 40 7. Estudios Molecular y Citogenético Molecular 40 7.1. Fijación 40 7.2. Protocolos de extracción de ADN genómico total 40-45 7.3. Purificación adicional del ADN genómico total 46 7.4. Medición de la concentración de ADN genómico total 46 7.5. Electroforesis en geles de agarosa 46-48 7.6. Digestión con endonucleasas de restricción 48-49 7.7. Extracción y purificación de fragmentos de ADN genómico de geles de agarosa 49-50 7.8. Marcaje de la sonda utilizando nick-translation 50 7.9. Hibridación in situ 50-53 8. Características y Condicionesde Endocría 53-54 RESULTADOS 55 9. Familia Coreidae 55 9.1. Complemento cromosómico y desarrollo meiótico de Athaumastus haematicus (Stål, 1859) 55-58 9.2. Complemento cromosómico y desarrollo meiótico de Hypselonotus bitriangulifer bitriangulifer Berg 58-59, 60 9.3. Complemento cromosómico y desarrollo meiótico de Phthia picta (Drury, 1770) 59-60 9.4. Complemento cromosómico y desarrollo meiótico de Leptoglossus impictus (Stål, 1859) 61-62 10. Familia Largidae 61 10.1. Complemento cromosómico y desarrollo meiótico de Largus fasciatus (Blanchard, 1843) 61, 63-65 10.2. Complemento cromosómico y desarrollo meiótico de Largus rufipennis Laporte, 1832 66-73 11. Familia Lygaeidae 73 11.1. Complemento cromosómico y desarrollo meiótico de Lygaeus alboornatus Blanchard, 1852 73-76 Citogenética Básica y Evolutiva de Heterópteros Fitófagos de Interés Agroeconómico de la Argentina III 12. Familia Pyrrhocoridae 77 12.1. Complemento cromosómico y desarrollo meiótico de Dysdercus albofasciatus Berg, 1878 77-81, 82 12.2. Complemento cromosómico y desarrollo meiótico de Dysdercus chaquensis Freiberg, 1948 81, 83-88 12.3. Complemento cromosómico y desarrollo meiótico de Dysdercus ruficollis (Linnaeus, 1764) 88, 89-91 12.4. Complemento cromosómico y desarrollo meiótico de Dysdercus imitator Blöte, 1931 92-96 13. Familia Rhopalidae 97 13.1. Complemento cromosómico y desarrollo meiótico de Jadera haematoloma (Herrich-Schaeffer, 1847) y Jadera sanguinolenta (Fabricius, 1775) 97-104 14. Estudios Moleculares y Citogenético-Moleculares 105 14.1. Comparación de los diferentes métodos de fijación 106, 107 14.2. Resultados de los distintos protocolos de extracción utilizados 106, 108 14.3. Resultados obtenidos de la digestión de las muestras de ADN genómico total con diferentes endonucleasas de restricción 109-113 14.4. Resultados obtenidos de la purificación adicional de las muestras de ADN genómico total 114 14.5 Hibridación in situ 114-117 DISCUSION 118-148 CONCLUSIONES GENERALES 149-153 ABREVIATURAS 154 SOLUCIONES 155-156 AGRADECIMIENTOS 157-158 REFERENCIAS 159-170 Citogenética Básica y Evolutiva de Heterópteros Fitófagos de Interés Agroeconómico de la Argentina IV RESUMEN El presente trabajo de Tesis se basa en el estudio citogenético básico y evolutivo de especies de Heteroptera fitófagas de interés agroeconómico para nuestro país. Se analizó el complemento cromosómico y el desarrollo meiótico de las siguientes especies: Hypselonotus bitriangulifer bitriangulifer Berg (2n= 19/20= 16+2m+X0/16+2m+XX), Athaumastus haematicus (Stål, 1859), Leptoglossus impictus (Stål, 1859) y Phthia picta (Drury, 1770) (las tres especies poseen 2n= 21/22= 18+2m+X0, 18+2m+XX) (Coreidae); Largus fasciatus (Blanchard, 1843) y Largus rufipennis Laporte, 1832 (ambas con 2n= 13/14= 12+X0/12+XX) (Largidae); Lygaeus alboornatus Blanchard, 1852 (2n= 12= 10+XY/10+XX) (Lygaeidae sensu lato); Dysdercus albofasciatus Berg, 1878 (2n= 12= 10+neoXY/10+neoXX), Dysdercus chaquensis Freiberg, 1948, Dysdercus imitator Blöte, 1931 y Dysdercus ruficollis (Linnaeus, 1764) (las tres especies con 2n= 13/14= 12+X0/12+XX) (Pyrrhocoridae); Jadera haematoloma (Herrich-Schaeffer, 1847) y Jadera sanguinolenta (Fabricius, 1775) (ambas con 2n= 13/14= 10+2m+X0/10+2m+XX) (Rhopalidae). En este trabajo se han encontrado alteraciones numéricas y estructurales en las especies analizadas de las familias Coreidae, Largidae, Lygaeidae y Pyrrhocoridae. Teniendo en cuenta el número cromosómico diploide modal para Coreidae, en Hypselonotus bitriangulifer bitriangulifer se observó una fusión autosómica que trajo aparejado una reducción del número de pares de autosomas. Los resultados obtenidos en Lygaeus alboornatus (Lygaeidae) muestran que esta especie posee el número cromosómico diploide más bajo mencionado hasta el presente para la subfamilia Lygaeinae, el cual se habría originado a partir del complemento cromosómico diploide atávico a través de una fusión autosómica. En la población de Largus fasciatus (Largidae) se encontró un individuo portador de un cromosoma B, mientras que en una población de Dysdercus albofasciatus (Pyrrhocoridae) cuatro especímenes presentaron un cromosoma supernumerario, que por sus características se habría originado a partir de un autosoma. En una población de Largus rufipennis (Largidae) se encontró un individuo portador de una translocación recíproca desigual entre dos cromosomas no homólogos de tamaño mediano. Finalmente, se encontraron individuos con alteraciones en los cromosomas sexuales en especímenes de Jadera sanguinolenta y J. haematoloma, los que presentaron un polimorfismo cromosómicoen relación con el tamaño del cromosoma sexual. Con respecto a los sistemas cromosómicos de determinación del sexo en Heteroptera, se encontró un sistema sexual neo-XY en Dysdercus albofasciatus, que es original para el género, así como para la familia Pyrrhocoridae, y para Heteroptera en virtud de su origen. Citogenética Básica y Evolutiva de Heterópteros Fitófagos de Interés Agroeconómico de la Argentina V En la meiosis de Heteroptera es característico que luego de paquitene siga el estadio difuso, el cual presenta diferencias entre las especies. En Dysdercus chaquensis (Pyrrhocoridae) se observó la ausencia de este estadio únicamente en especímenes pertenecientes a una población endocriada, mientras que estaba presente en poblaciones naturales. La aplicación de la técnica de impregnación argéntica a células meióticas de Dsydercus imitator (Pyrrhocoridae) permitió observar la presencia de varios cuerpos prenucleolares a lo largo de toda la meiosis. El ciclo peculiar del nucléolo pudo relacionarse con características del ciclo de vida de la especie. Se realizó la extracción de ADN genómico total a partir de ejemplares enteros de Athaumastus haematicus, Hypselonotus bitriangulifer bitriangulifer, Leptoglossus impictus, Phthia picta (Coreidae), Largus fasciatus, Largus rufipennis (Largidae), Lygaeus alboornatus (Lygaeidae), Dysdercus albofasciatus, Dysdercus chaquensis, Dysdercus imitator, Dysdercus ruficollis (Pyrrhocoridae) y Jadera sanguinolenta (Rhopalidae), mediante cuatro métodos distintos de extracción. El análisis comparativo de los resultados obtenidos con cada uno de ellos mostró que
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