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La Disploidía Como Motor Evolutivo En La Familia Asteraceae Alicia

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La Disploidía Como Motor Evolutivo En La Familia Asteraceae Alicia Trabajo de Final de Grado Grado en Biología La disploidía como motor evolutivo en la familia Asteraceae Aportación de datos cariológicos en la alianza de las helianteas Alicia Paniego Gómez Tutora externa: Sònia Garcia Giménez Tutor interno: Joan Vallès Xirau Lugares de realización del trabajo: Facultad de Farmacia y Ciencias de la Alimentación de la Universitat de Barcelona, Centros Científicos y Tecnológicos de la Universitat de Barcelona e Instituto Botánico de Barcelona Julio de 2017 Fotografía de la cubierta: Lindheimera texana A.Gray & Engelm. http://basia.typepad.com/india_ink/2010/05/texas-wildflowers-medley.html. Lady Bird Johnson Wildflower Center, Austin, TX. Índice 1. Introducción ................................................................................................................. 1 1.1. Familia Asteraceae ......................................................................................................... 1 1.2. La alianza de las helianteas .......................................................................................... 2 1.3. Recuentos cromosómicos ............................................................................................. 4 1.4 Estimación de la cantidad de ADN nuclear ................................................................ 5 2. Objetivos ...................................................................................................................... 6 3. Materiales y metodología ........................................................................................... 7 3.1. Siembra de aquenios en placa ..................................................................................... 7 3.2. Siembra de aquenios en tiesto ..................................................................................... 7 3.3. Conteo de cromosomas ................................................................................................. 8 3.4. Estimación de la cantidad de ADN nuclear por citometría de flujo ....................... 10 4. Resultados ................................................................................................................. 12 5. Discusión .................................................................................................................... 19 5.1. Datos obtenidos para las especies de la alianza de las helianteas...................... 19 5.2. Datos obtenidos para el resto de especies .............................................................. 27 6. Conclusiones ............................................................................................................. 28 7. Agradecimientos ....................................................................................................... 29 8. Bibliografía ................................................................................................................. 31 Summary With this final degree project we aim to contribute new data on genome sizes and chromosome numbers for a group of species belonging to family Asteraceae, in particular from subfamily Asteroideae. This group is known as the Heliantheae alliance, an assembly of ten tribes without any taxonomic rank. In order to study the chromosome numbers, the technique used has been the classical squash method with orcein staining. Nuclear genome size has been assessed by flow cytometry with propidium iodide as a fluorochrome. We have obtained new information for 13 species, seven genera and two tribes of the family, of which ten species, six genera and also two tribes belong to the Heliantheae alliance. Our data contribute to the pool of knowledge on genome size and chromosome numbers in this alliance (and other Asteraceae taxa) and will further allow deeper studies and a better understanding on the role of dysploidy in the evolution of this specific group. However, we still lack data for tribes Chaenactidae, Neurolaeneae and Polymnieae to complete the alliance representation. 1 1. Introducción 1.1. Familia Asteraceae La familia Asteraceae, también conocida como Compositae, es la que posee mayor diversidad y riqueza dentro de las angiospermas. La mayoría de las especies que la forman son herbáceas y no suelen ser suculentas. Pueden ser anuales, bienales o perennes y existen especies hidrofíticas, heliofíticas, mesofíticas y xerofíticas. Como es lógico en la mayor familia de plantas en número de especies, se trata de un grupo de distribución muy cosmopolita (Jeffrey, 2007). Las flores se disponen en una inflorescencia que recibe el nombre de capítulo. En esta, el pedúnculo se ensancha formando el receptáculo, rodeado por el involucro que es donde se disponen las flores. Hay tres tipos de capítulos: 1. Con flores tubulares en el centro (que suelen ser hermafroditas) y liguladas en la periferia (por lo general, femeninas) (Figura 1). 2. Solo con flores tubulares, hermafroditas. 3. Solo con flores liguladas, hermafroditas. Figura 1. Inflorescencia en capítulo. Flores liguladas externas y tubulares internas Dirección URL: <http://basica.primariatic.sep.gob.mx/descargas/colecciones/proyectos/red_escolar/publi_reinos/flora/mar garita/galeria.htm>. [Consulta: 29/3/2017]. 1 La familia Asteraceae es un grupo monofilético que consta de numerosas tribus, que se distribuyen en 12 subfamilias (Funk et al., 2009): -Subfamilia Barnadesioideae, que consta de una tribu, Barnadesieae. -Subfamilia Mutisioideae, cuyos géneros se agrupan en las tribus Onoserideae, Mutisieae y Nassauvieae. -Subfamilia Stifftioideae, que solo tiene una tribu llamada Stifftieae. -Subfamilia Wunderlichioideae, con dos tribus: Wundelichieae y Hyalideae. -Subfamilia Gochnatioideae, con una sola tribu denominada Gochnatieae. -Subfamilia Hecastocleidoideae, que posee también una tribu, Hecastoclideae. -Subfamilia Carduoideae, con tres tribus: Cynareae, Dicomeae y Tarchonantheae. -Subfamilia Pertyoideae, cuyas especies se incluyen en la tribu Pertyeae. -Subfamilia Gymnarrhenoideae, compuesta por la tribu Gymnarrheneae. -Subfamilia Cichorioideae, que posee las siguientes tribus: Gundelieae, Cichorieae, Arctotideae, Liabeae y Vernonieae. -Subfamilia Corymbioideae, con la tribu Corymbieae. -Subfamilia Asteroideae, la de mayor riqueza de las Asteraceae y que recoge 20 tribus: Anthemideae, Astereae, Athroismeae, Bahieae, Calenduleae, Chaenactideae, Coreopsideae, Eupatorieae, Feddeae, Gnaphalieae, Heleniae, Heliantheae, Inuleae, Madieae, Millerieae, Neurolaeneae, Perityleae, Polymnieae, Senecioneae y Tageteae (Figura 2). 1.2. La alianza de las helianteas La alianza de las helianteas es una agrupación sin rango taxonómico de 10 tribus de la subfamilia Asteroideae; posee aproximadamente 5500 especies, que constituyen el 20-30% de la familia Asteraceae. Las tribus que pertenecen a esta alianza son: Bahiae, Chaenactidae, Eupatorieae, Heleniae (constituida a su vez por las subtribus Gaillardiinae, Tetraneurinae, Psathyrotinae y Marshalliinae), Heliantheae, Madieae, Millerieae, Neurolaeneae, Polymnieae y Tageteae (Funk et al., 2009) (Figura 2). 2 Figura 2. Fragmento de un árbol filogenético de las Asteraceae que incluye la subfamilia Asteroideae (Funk et al., 2009). Algunos caracteres morfológicos de las tribus que pertenecen a la alianza de las helianteas son: la presencia de brácteas seriadas, el ennegrecimiento de las anteras, estilos con pelos y apéndices más cortos que los estigmas (excepto en la tribu Eupatorieae) y la presencia de una capa de fitomelanina en el aquenio. Se trata de un grupo bastante reciente desde el punto de vista evolutivo. La alta variedad morfológica que presentan sus especies explica la dificultad que ha habido históricamente para hacer una clasificación práctica de éstas (Funk et al., 2009). La mayoría de las especies pertenecientes a la alianza de las helianteas son endémicas de América del Norte (Figura 2). Se considera que la disploidía (aumento o descenso del número de base de cromosomas, generalmente mediante procesos de fusión o fisión) ha jugado un papel fundamental en la evolución de este grupo (Mota et al., 2016). En las tribus que componen la alianza se ha podido observar un incremento del número básico de cromosomas, mediante procesos de disploidía, hasta x = 19, llegando a ser más del doble del número básico más frecuente en la familia (x = 9). La más representativa es la tribu Heliantheae (que da nombre al grupo), con más de 5200 especies (Jeffrey, 2007). 3 1.3. Recuentos cromosómicos Antes de que se desarrollasen estudios filogenéticos basados en el análisis de las secuencias de ADN, el análisis del número de cromosomas desde los puntos de vista filogenético y evolutivo se veía obstaculizado por la falta de información que se tenía sobre qué géneros o tribus eran basales (las más antiguas del grupo). Sin embargo, con la introducción de dichas técnicas se empezaron a comparar los datos moleculares con los datos del número de cromosomas para averiguar cómo trazar dicha evolución (Semple & Watanabe, 2009). En la actualidad, muchos de los datos de número de cromosomas se encuentran bases de datos accesibles por internet, como por ejemplo el “Index to Plant Chromosome Numbers” (http://www.tropicos.org/Project/IPCN), un proyecto del Jardín Botánico de Missouri; la “Chromosome Counts Database” (http://ccdb.tau.ac.il/), organizada por investigadores de la Universidad de Tel Aviv; o la base de datos “Index to Chromosome Numbers in Asteraceae” (http://www.lib.kobe-u.ac.jp/infolib/meta_pub/engG0000003asteraceae), construida por el Dr. K. Watanabe de la Universidad de Kobe. Los primeros recuentos de cromosomas
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