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Determinación Sexual En Primates Neotropicales: El Caso De Los Monos Aulladores

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Determinación Sexual En Primates Neotropicales: El Caso De Los Monos Aulladores Tesis Doctoral Determinación sexual en primates neotropicales: el caso de los monos aulladores Steinberg, Eliana Ruth 2011 Este documento forma parte de la colección de tesis doctorales y de maestría de la Biblioteca Central Dr. Luis Federico Leloir, disponible en digital.bl.fcen.uba.ar. Su utilización debe ser acompañada por la cita bibliográfica con reconocimiento de la fuente. This document is part of the doctoral theses collection of the Central Library Dr. Luis Federico Leloir, available in digital.bl.fcen.uba.ar. It should be used accompanied by the corresponding citation acknowledging the source. Cita tipo APA: Steinberg, Eliana Ruth. (2011). Determinación sexual en primates neotropicales: el caso de los monos aulladores. Facultad de Ciencias Exactas y Naturales. Universidad de Buenos Aires. Cita tipo Chicago: Steinberg, Eliana Ruth. "Determinación sexual en primates neotropicales: el caso de los monos aulladores". Facultad de Ciencias Exactas y Naturales. Universidad de Buenos Aires. 2011. Dirección: Biblioteca Central Dr. Luis F. Leloir, Facultad de Ciencias Exactas y Naturales, Universidad de Buenos Aires. Contacto: [email protected] Intendente Güiraldes 2160 - C1428EGA - Tel. (++54 +11) 4789-9293 UNIVERSIDAD DE BUENOS AIRES FACULTAD DE CIENCIAS EXACTAS Y NATURALES DEPARTAMENTO DE ECOLOGÍ A, GENÉTI CA Y EVOLUCI ÓN “Determinación sexual en Primates Neotropicales: el caso de los monos aulladores” Tesis presentada para optar al título de Doctor de la Universidad de Buenos Aires en el área Ciencias Biológicas Lic. Eliana Ruth Steinberg Directora de Tesis: Prof. Dra. Marta Dolores Mudry Consejera de Estudios: Prof. Dra. Marta Dolores Mudry Lugar de trabajo: Grupo de Investigación en Biología Evolutiva (GIBE), Departamento de Ecología, Genética y Evolución, Facultad de Ciencias Exactas y Naturales, Universidad de Buenos Aires Buenos Aires, Argentina Marzo, 2011 Determinación sexual en Primates Neotropicales: el caso de los monos aulladores La Cariosistemática perm ite comparar taxa relacionados, en particular especies. Una variable de análisis con valor diagnóstico es el tipo de sistema de determinación sexual. En este Trabajo de Tesis se analizaron monos aulladores del género Alouatta con sistemas de determinación sexual múltiple resultado de translocaciones Y-autosoma. Se los comparó con otros 3 Ceboidea (Cebus, Saimiri y Aotus) y con dos Hominoidea (Pan troglodytes y Homo sapiens) ante la diversidad de patrones de determinación sexual, en particular en machos. Se realizó la primera caracterización del cariotipo de Alouatta pigra que mostró 2N= 58, X1X1X2X2 / X1X2Y1Y2. Se estudió conservación genómica por FISH con las sintenias 3/15 y 3/21 evidenciando que estas asociaciones no estarían conservadas en las especies mesoamericanas A. pigra y A. palliata. Este estudio junto al de homeologías cromosómicas, mostró que los autosomas involucrados en las translocaciones que darían origen a los multivalentes sexuales en las especies sudamericanas y mesoamericanas serían distintos. En el marco conceptual de “Evidencia Total”, el análisis combinado de variables moleculares y cromosómicas resolvió las relaciones de parentesco entre las especies de aulladores de ambos orígenes americanos, demostrando su utilidad en el esclarecimiento de controversias que relacionan la Taxonomía y la Evolución de los primates ceboideos. Palabras claves: Alouatta, evolución cromosómica, Meiosis, sistemas de determinación sexual, Evidencia Total, SRY, sintenias cromosómicas, filogenias por análisis combinado. Sex determination in neotropical primates: the example of the Howler Monkeys Karyosystematics allows for comparison of related taxa, particularly species. The sex determination system constitutes a variable with diagnostic value. In this dissertation howler monkeys, genus Alouatta, with multiple sex determination systems product of Y-autosome translocations, were analized. In light of the diversity of sex determination patterns observed, particularly in males, they were compared with other 3 Ceboidea (Cebus, Saim iri and Aotus) and with 2 Hominoidea (Pan troglodytes and Hom o sapiens). The first cytogenetic characterization of Alouatta pigra karyotype was performed, exhibiting a 2N= 58, X1X1X2X2/X1X2Y1Y2. Genom ic conservation was studied by FISH analizing the syntenies 3/15 and 3/21, showing that these asociations are not conserved in the mesoamerican howlers, A. pigra y A. palliata. This analysis, together with the chromosomal homologies, demonstrated that the autosomal pairs involved in the translocations that originate the sexual multivalents are different in the southamerican and mesoamerican species. Under the framework of the “Total Evidence” concept, the combined análisis of molecular and chromosomal variables resolved the phylogenetic relationships between the howler species of both american origins, demonstrating its utility to clarify the controversies related to the Taxonomy and Evolution of Ceboidea primates. Key words: Alouatta, chromosome evolution, Meiosis, sex determination systems, Total Evidence, SRY, chromosomal syntenies, phylogenies by combined analysis. Agradecimientos A la Universidad de Buenos Aires (UBACyT X107 y X145 M.D.M.), en particular a la Facultad de Ciencias Exactas y Naturales y al departamento de Ecología, Genética y Evolución que posibilitaron y facilitaron mi formación doctoral así cómo el desarrollo experimental objeto de este trabajo de Tesis. Al CONICET, institución nacional que mediante los subsidios PIP 5012 y 112-200801-00744 concedidos a mi Directora de Tesis y las becas internas de postgrado tipo I y II que me fueron otorgadas avaló la concreción de este trabajo de Tesis Doctoral. A los directores de las Instituciones, tales como Jardines Zoológicos, Bioterios y centros de cría, que permitieron la realización estos estudios así como a los profesionales, veterinarios y cuidadores, que colaboraron en la obtención de las muestras y seguimiento de los anim ales de cautiverio objeto de estudio. A la Dra. Marta D. Mudry, mi directora de Tesis, mi maestra en la ciencia y en la vida. Gracias por enseñarme a amar la Biología y a siempre tratar de superarnos. A la Dra. Mariela Nieves por su invaluable colaboración en el desarrollo e interpretación de las técnicas citogenéticas. Al Lic. Alejandro Laudicina, por su paciencia para responder a mis infinitas preguntas sobre FISH y ayudarme a sacar adelante la técnica en los ceboideos. A la Dra. M. García Caldés por su contribución a la discusión de las homeologías en aulladores. Al equipo de investigadores de la Unitat de Biologia Cellular i Genètica Mèdica, Facultat de Medicina, de la Universidad Autónoma de Barcelona por su colaboración en el análisis por inmunofluorescencia de células germinales. Al equipo mexicano de investigadores y a todo el equipo de campo por su colaboración en la obtención de las muestras de aulladores de distribución natural marginal norte. A Juan Pablo Luaces y Luis Rossi, por hacer de mi estadía en FMed una grata experiencia y por el intercambio de saberes en la aplicación e interpretación de las técnicas citogenéticas. A los jurados que evaluarán mi trabajo de Tesis Doctoral, que con sus aportes y sugerencias ayudarán a mejorarlo. A Gaby, Mae, Lucía, Nancy, Carola, Gise, Ana, Cristina y Leandro por compartir conmigo el camino de la ciencia con mates, aguante y continuas charlas. A mis amigos, por comprenderme, soportarme y acompañarme durante los años que duró mi doctorado. A Carlos, por estar a mi lado, por comprenderme, por su paciencia y por ser mi cable a tierra durante todos estos años. A mis padres y a mi hermana, por el apoyo incondicional y por hacerme la persona que soy hoy. Í ndice 1. I ntroducción……………………………………………………………………………………………………… 1 1.1 Rearreglos cromosómicos y especiación………………………………………………………… 1 1.2 Cariosistem ática………………………………………………………………………………………………. 2 1.2.1 Cromosomas sexuales, especiación y cariosistem ática………………………… … 3 1.2.2 Mecanismos de determinación sexual……………………………………………………... 4 1.2.2.1 Plantas……………………………………………………………………………………………………. 5 1.2.2.2 I nvertebrados……………………………………………………………………………………….. 7 1.2.2.3 Vertebrados………………………………………………………………………………………….. 10 1.2.2.3.1 Prim ates……………………………………………………………………………………………. 14 1.2.3 Sintenias crom osóm icas…………………………………………………………………………….. 17 1.2.3 1 Prim ates……………………………………………………………………………………………….. 19 1.2.4 El gen SRY…………………………………………………………………………………………………. 22 1.2.4.1 Prim ates…………………………………………………………………………………………………. 23 1.3 El m odelo de estudio………………………………………………………………………………………. 24 1.3.1 El género Alouatta…………………………………………………………………………………….. 26 1.3.1.1 Diagnosis citogenética………………………………………………………………………… 29 1.3.1.2 Asociaciones cromosómicas y sistemas de determinación sexual……… 31 1.3.1.3 Estudios filogenéticos…………………………………………………………………………. 32 2. Objetivos e Hipótesis………………………………………………………………………………………. 34 2.1 Objetivos…………………………………………………………………………………………………......... 34 2.1.1 Objetivo General………………………………………………………………………………………… 34 2.1.2 Objetivos específicos…………………………………………………………………………………. 34 2.2 Hipótesis de trabajo………………………………………………………………………………………… 3 3. Materiales y Métodos………………………………………………………………………………………… 35 3.1 Material biológico…………………………………………………………………………………………….. 35 3.2 Diagnosis por caracterización citogenética clásica…………………………………………. 38 3.2.1 Estudios en células somáticas…………………………………………………………………… 38 3.2.1.1 Extracción de las muestras………………………………………………………………….. 38 3.2.1.2 Cultivo de sangre periférica…………………………………………………………………. 38 3.2.1.3 Análisis de metafases……………………………………………………………………………
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