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TESIS CODIUM.Pdf UNIVERSIDAD NACIONAL DEL SUR TESIS DOCTOR EN BIOLOGÍA BIOLOGÍA Y ULTRAESTRUCTURA DE CODIUM SPP. (BRYOPSIDOPHYCEAE, CHLOROPHYTA): MORFOLOGÍAS VEGETATIVA Y REPRODUCTIVA, CICLOS DE VIDA Y EPIFITISMO ALICIA BEATRIZ MIRAVALLES BAHIA BLANCA ARGENTINA 2008 UNIVERSIDAD NACIONAL DEL SUR TESIS DOCTOR EN BIOLOGÍA BIOLOGÍA Y ULTRAESTRUCTURA DE CODIUM SPP. (BRYOPSIDOPHYCEAE, CHLOROPHYTA): MORFOLOGÍAS VEGETATIVA Y REPRODUCTIVA, CICLOS DE VIDA Y EPIFITISMO ALICIA BEATRIZ MIRAVALLES BAHIA BLANCA ARGENTINA 2008 PREFACIO Esta Tesis se presenta como parte de los requisitos para optar al grado Académico de Doctor en Biología, de la Universidad Nacional del Sur y no ha sido presentada previamente para la obtención de otro título en esta Universidad u otra. La misma contiene los resultados obtenidos en investigaciones llevadas a cabo en el ámbito del Departamento de Biología, Bioquímica y Farmacia durante el período comprendido entre el 07/09/99 y el 09/12/08, bajo la dirección del Doctor Eduardo J. Cáceres, Profesor Titular de Biología de Algas y Hongos e Investigador Principal de la Comisión de Investigaciones Científicas de la Provincia de Buenos Aires (CIC) y de la Doctora Patricia I. Leonardi, Profesora Adjunta de Biología de Algas y Hongos e Investigadora Independiente del Consejo Nacional de Investigaciones Científicas y Técnicas (CONICET). 09 de diciembre de 2008 Departamento de Biología, Bioquímica y Farmacia Universidad Nacional del Sur UNIVERSIDAD NACIONAL DEL SUR Secretaría General de Posgrado y Educación Continua La presente tesis ha sido aprobada el .…/.…/.….. , mereciendo la calificación de ......(……………………) AGRADECIMIENTOS Durante la realización de esta tesis muchas personas me brindaron su ayuda, su colaboración, sus críticas y principalmente su apoyo necesarios para su culminación: A los Doctores Eduardo J. Cáceres y Patricia I. Leonardi por guiarme y acompañarme en todo este proceso, por brindarme toda su experiencia y contagiarme su entusiasmo. Al Departamento de Biología, Bioquímica y Farmacia, lugar donde se realizaron los estudios. A mis amigos del Laboratorio de Ficología y Micología: Patricia Leonardi, Virginia Bianchinotti, Karina Michetti, Nora Ferrer, Cecilia Damiani, Amelia Vouilloud, Cecilia Popovich, Romina Sánchez y Lucas Martín, por su apoyo incondicional, por todos los momentos compartidos… A Pablo Martín, Lucas Martín, Sandra Fiori, Sebastián Bronfen, quienes me trajeron muestras de Codium de sus viajes de muestreo. Al personal de Microscopía Electrónica del CRIBABB: Viviana Sorrivas, M. Julia Yañez, Carina Cano y Carlos Jones. A mis amigas Andrea Tomás y Marina Díaz. A mis compañeros de Farmacobotánica e Histología y Biología Celular. A mis padres y hermana, que en todo momento me apoyaron para que pueda concretar mis metas. A Javier, Victoria y Antonio, que con todo su amor, paciencia, comprensión y constante estímulo me alentaron para que pueda concluir esta etapa. Con la alegría de llegar a la recta final, deseo redundar en mi agradecimiento a todas y cada una de las personas que de una forma u otra han ayudado a que esto fuera realidad. ii RESUMEN Se estudiaron poblaciones de tres taxa del género Codium que crecen en las costas de las Provincias de Buenos Aires, Río Negro y Chubut: C. decorticatum, C. fragile subsp. novae-zelandiae y C. vermilara. Los talos de los tres taxa, erguidos y ramificados dicotómicamente, se fijaban al sustrato mediante un disco basal. El tamaño y forma de los utrículos y gametangios, presencia/ausencia de mucrón fueron las características principales que permitieron diferenciar los taxa. Ultraestructuralmente los utrículos mostraron una vacuola central rodeada por una fina capa de citoplasma parietal en la que se encontraban las organelas. La ultraestructura de los pelos y de los filamentos medulares fue similar a la de los utrículos. Los gametangios se formaban a los lados de los utrículos. Se observaron gametangios femeninos en los tres taxa y masculinos en C. fragile. Los masculinos eran más delgados que los femeninos. La liberación de ambos tipos de gametas inmersas en abundante mucílago se produjo a través de la ruptura del extremo apical. Este es el primer estudio ultraestructural del proceso de gametogénesis y de las gametas femeninas y masculinas en el género Codium. Las gametogénesis se iniciaban con la división de núcleos y cloroplastos. La delimitación de las gametas se producía por la actividad de vesículas dictiosómicas que rodeaban porciones de protoplasma uninucleadas. iii Las gametas femeninas eran piriformes; poseían una papila apical de la que emergían los flagelos. Anteriormente se encontraban el núcleo y una mitocondria grande con forma de V invertida. El resto de la gameta estaba ocupado por numerosos cloroplastos. Las gametas masculinas eran ovoides alargadas, de menor tamaño; poseían dos o tres cloroplastos escasamente desarrollados y una papila apical hialina de la que emergían los flagelos. Se comprobó la germinación partenogenética de las gametas femeninas de C. fragile en ausencia de gametas masculinas. También se observó la germinación intragametangial de las gametas femeninas. Se verificó la reproducción sexual en C. fragile en presencia de ambos tipos de gametas por la presencia de cigotos tetraflagelados y su posterior germinación. Para cada uno de los tres taxa se proporcionó información cariológica, con un número haploide de 10, como el obtenido por otros autores para otras especies del género. Se confirmó la partenogénesis para las tres especies. Se determinó que C. vermilara y C. decorticatum presentaban un ciclo de vida haplobionte haplonte y C. fragile un ciclo de vida haplobionte haplonte o diplonte, de acuerdo a la capacidad de formación de gametas masculinas. Se identificaron las distintas especies de epífitos sobre C. vermilara de la localidad de Las Grutas. Los géneros de epífitos más comunes fueron Ceramium, Polysiphonia, Callithamnion y Ectocarpus. De acuerdo al grado de invasión de las estructuras de fijación se diferenciaron cuatro modos de fijación de los epífitos al hospedante. Se estudió con microscopía óptica y electrónica la interfase entre los epífitos y el hospedante para determinar el grado de daño que iv ejercía el epífito al hospedante. Las distintas especies de epífitos que crecían sobre Codium y entre sus utrículos no causaban daño al talo. v ABSTRACT The purpose of this thesis was to study populations of C. decorticatum, C. fragile subsp. novae-zelandiae and C. vermilara of the genus Codium growing on the coastal areas of the provinces of Buenos Aires, Río Negro and Chubut, Argentina. Their thalli were found to be erect and dichotomously branched and they were fixed to the substrate by a basal disk. The following characteristics were crucial for the differentiation of these taxa: i) size and shape of utricula and gametangia, and ii) presence and/or absence of mucron. At the ultrastructural level, it could be observed that utricula showed a central vacuole which was surrounded by a thin layer of parietal cytoplasm in which organella were located. The ultrastructure of hairs and medular filaments was similar to that of utricula. Gametangia were formed on the sides of utricula. Female gametangia were observed in the three taxa analyzed whereas male gametangia were found only in C. fragile and they were thinner than the female ones. The release of both types of gametes, which were immersed in abundant mucilage, occurred through a slit. This is the first ultrastructural study conducted to date on gametogenesis and on female and male gametes in the genus Codium. Findings from this thesis indicate that gametogenesis begins with the division of nuclei and chloroplasts and that gametes are individualized as a result of the activity of dyctiosomic vesicles which surround uninucleate portions of protoplasm. It could also be vi observed that female gametes are pyriform and that they have an apical papilla from which flagella emerge. The nucleus and a large inverted V-shaped mitochondrium were found to be located in the anterior region of gamete while the rest of the gamete was occupied by numerous chloroplasts. Male gametes were found to be ovoid and elongated, they were smaller than female gametes and they had two or three chloroplasts poorly developed and a hyaline apical papilla from which flagella emerged. Parthenogenetic germination of female gametes of C. fragile was observed in the absence of male gametes. Intragametangial germination of female gametes was also observed. On the other hand, the presence of tetraflagellate zygotes and their subsequent germination in C. fragile was indicative of sexual reproduction in this species. Karyological information was provided for each of the three taxa with a haploid number of 10. The latter agreed with that obtained by other authors for other species of the genus. Parthenogenesis was confirmed for the three species. Also, it could be concluded that C. vermilara and C. decorticatum have a haplobiont haplont life cycle and that C. fragile, depending on the presence or absence of male gametes, has either a haplobiont diplont or haplont life cycle. Different species of epiphytes on C. vermilara could also be identified in Las Grutas, Argentina, the most common epiphytic genera being Ceramium, Polysiphonia, Callithamnion and Ectocarpus. Taking into account the degree of invasion of fixing structures, four modes of attachment of epiphytes to the host were identified. The epiphyte-host interface was studied via optical and electron microscopy in order to determine the degree of damage exerted
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