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Chloridoideae- Poaceae) UNIVERSIDAD NACIONAL DEL LITORAL Facultad de Bioquímica y Ciencias Biológicas Tesis para la obtención del Grado Académico de Doctor en Ciencias Biológicas EVOLUCIÓN, DESARROLLO Y ESTRUCTURA DE LAS INFLORESCENCIAS EN LA SUBTRIBU ELEUSININAE (CHLORIDOIDEAE- POACEAE) Sebastián E. Muchut Director de Tesis: Abelardo C. Vegetti Co-director de Tesis: Renata Reinheimer Cátedra de Morfología Vegetal Departamento de Biología Vegetal Facultad de Ciencias Agrarias – Universidad Nacional del Litoral -2017- Agradecimientos Quiero expresar mi agradecimiento a la Facultad de Bioquímica y Ciencias Biológicas de la Universidad Nacional del Litoral (UNL) por permitirme realizar el “Doctorado en Ciencias Biológicas”, especialmente al personal de la oficina de posgrado por resolver siempre atentamente mis inquietudes. A la Facultad de Ciencias Agrarias (FCA-UNL) por brindarme todas las instalaciones y herramientas necesarias para la realización de esta tesis. A la Agencia Nacional de Promoción Científica y Tecnológica (ANPCyT) y al Consejo Nacional de Investigaciones Científicas y Técnicas (CONICET), por haberme otorgado las becas de posgrado que me permitieron dedicarme exclusivamente a la tesis doctoral. A mis directores de tesis, los Dres. Abelardo Vegetti y Renata Reinheimer por darme la oportunidad de iniciarme en la investigación científica, por enseñarme todo lo referido a la morfología, taxonomía, filogenia y evolución de gramíneas, por sus instrucciones con respecto a la escritura científica, por sus consejos, paciencia, y por todo el apoyo brindado durante estos años. A Abelardo, por haberme dado la posibilidad de integrar su grupo de trabajo, transmitirme su pasión por la investigación, aconsejarme constantemente e instruirme sobre el mundo científico. A la cátedra de Morfología Vegetal, por integrarme al grupo desde el primer día y hacerme sentir uno más de ellos, por los momentos académicos y recreativos compartidos. A Juan Carlos Tivano, Vanesa Pilatti, Pablo Ortega y su equipo, por su predisposición y asistencia en los viajes de colección que me permitieron encontrar las plantas para hacer los estudios de desarrollo. A Vanesa Pilatti, Nora Uberti-Manassero y Andrea Reutemann, por ser mi grupo de apoyo y guía durante toda la tesis, por enseñarme todas las técnicas, por los intensos debates científicos (y no tan científicos), consejos académicos (y no tan académicos) y, especialmente, por la buena onda y amistad que me brindaron estos años. A mi familia, por apoyarme incondicionalmente y alentarme en todas mis decisiones. A Andre, mi compañera en esta vida, por tu amor y aliento diario. Por entender mi pasión por este trabajo y empujarme siempre a ser mejor persona y profesional. PPuubblliiccaacciioonneess Muchut, S. E., Pilatti, V., Uberti-Manassero, N., Vegetti, A. C., & Reinheimer, R. (2017). Inflorescence diversity in subtribe Eleusininae (Poaceae: Chloridoideae: Cynodonteae). Flora, 228: 50-59. Muchut, S. E., Pilatti, V., Uberti-Manassero, N. G., Perreta, M., Vegetti, A. C. (2017). Trophotagma enrichment axes in Poaceae. Botanical Review, 1-13 Muchut, S. E., Reutemann, A. G., Uberti Manassero, N. G., Vegetti, A. C. (2017). Synflorescence morphology of grasses with reduced terminal inflorescences: a case study of Jouvea (Cynodonteae, Chloridoideae, Poaceae). Phytotaxa, 302: 241-250. “…el ‘espectáculo’ de la evolución sería una serie de acontecimientos, asombrosamente improbable, suficientemente perceptible cuando es observado en retrospectiva, y sujeto a explicación rigurosa, pero absolutamente impredecible y relativamente irrepetible. Hagamos retroceder la cinta magnetofónica de la vida hasta los primeros días de Burgess Shale; hagámosla sonar de nuevo desde un punto de partida idéntico, y tendremos una posibilidad tan reducida que es casi inexistente de que algo parecido a la inteligencia humana adorne la melodía que se escuchará.” Stephen Jay Gould, La vida maravillosa ÍNDICE Abreviaturas y símbolos ................................................................................................. 3 Resumen ....................................................................................................................... 4 Abstract ......................................................................................................................... 5 INTRODUCCIÓN ........................................................................................................... 6 La familia Poaceae (Gramíneas) ................................................................................ 7 Las espiguillas e inflorescencias de gramíneas .......................................................... 8 Evolución de la morfología ....................................................................................... 11 Estudios de desarrollo de inflorescencias ................................................................. 12 La subtribu Eleusininae (Cynodonteae-Chloridoideae) ............................................. 12 Hipótesis de trabajo .................................................................................................. 14 OBJETIVOS DEL TRABAJO ...................................................................................... 15 Objetivo general ....................................................................................................... 16 Objetivos específicos ............................................................................................... 16 MATERIALES Y MÉTODOS ........................................................................................ 17 Morfología adulta y diversidad de inflorescencias..................................................... 18 Evolución de inflorescencias .................................................................................... 18 Extracción de ADN, amplificación y secuenciación ............................................... 19 Reconstrucción filogenética .................................................................................. 19 Codificación de datos morfológicos ....................................................................... 20 Estimación de estados ancestrales ....................................................................... 20 Tasas de transición y Modelos de evolución ......................................................... 21 Coevolución de caracteres morfológicos .............................................................. 21 Evolución condicional– Orden de adquisición de estado de carácter .................... 22 Análisis ontogénico .................................................................................................. 23 RESULTADOS ............................................................................................................ 24 Morfología de inflorescencia ..................................................................................... 25 Tipos de inflorescencia ......................................................................................... 27 Análisis evolutivos .................................................................................................... 29 Reconstrucción ancestral ...................................................................................... 29 Tasas de transición y Modelos de evolución ......................................................... 37 Correlación evolutiva entre caracteres de inflorescencia ...................................... 37 Desarrollo de inflorescencias y espiguillas ............................................................... 46 Transición de meristema vegetativo a meristema de inflorescencia ...................... 46 Sentido de iniciación de las ramas primarias sobre el eje principal ....................... 46 Interrupción del desarrollo del meristema de inflorescencia .................................. 46 Sentido de diferenciación de las ramas primarias sobre el eje principal ................ 46 Sentido de iniciación de las ramas secundarias sobre las ramas primarias .......... 47 Sentido de diferenciación de las ramas secundarias sobre las ramas primarias ... 47 Desarrollo de las espiguillas ................................................................................. 55 Diferenciación de las espiguillas ........................................................................... 55 Filotaxis de antecios ............................................................................................. 58 DISCUSIÓN DE RESULTADOS Y CONCLUSIONES ................................................. 59 Inflorescencias de la subtribu Eleusininae ................................................................ 60 Evolución de las inflorescencias ............................................................................... 61 Aportes de los estudios de desarrollo ....................................................................... 65 Conclusiones ............................................................................................................ 69 BIBLIOGRAFÍA ............................................................................................................ 71 APÉNDICES ................................................................................................................ 84 2 AAbbrreevviiaattuurraass yy ssíímmbboollooss Br: bráctea Me: meristema de espiguilla ca: carpelo Mf: meristema floral e: estambre Mf1: primer meristema floral g: gineceo Mf2: segundo meristema floral Gi: gluma inferior Mf3: tercer meristema floral Gs: gluma superior MI: meristema de inflorescencia Hv: hoja
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