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Comunicación Química De Anuros: Una Tesis Doctoral Comunicación química de anuros: una perspectiva integral a partir de aspectos comportamentales, morfológicos y químicos en dos especies de Hypsiboas (Amphibia: Anura: Hylidae) Brunetti, Andrés E. 2014-12-19 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: Brunetti, Andrés E.. (2014-12-19). Comunicación química de anuros: una perspectiva integral a partir de aspectos comportamentales, morfológicos y químicos en dos especies de Hypsiboas (Amphibia: Anura: Hylidae). Facultad de Ciencias Exactas y Naturales. Universidad de Buenos Aires. Cita tipo Chicago: Brunetti, Andrés E.. "Comunicación química de anuros: una perspectiva integral a partir de aspectos comportamentales, morfológicos y químicos en dos especies de Hypsiboas (Amphibia: Anura: Hylidae)". Facultad de Ciencias Exactas y Naturales. Universidad de Buenos Aires. 2014-12-19. 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 Biodiversidad y Biología Experimental Comunicación química en anuros: una perspectiva integral a partir de aspectos comportamentales, morfológicos y químicos en dos especies de Hypsiboas (Amphibia: Anura: Hylidae) Tesis presentada para optar por el título de Doctor de la Universidad de Buenos Aires en el área de Ciencias Biológicas Andrés E. Brunetti Directores de tesis: Julián Faivovich y Gabriela M. Cabrera Consejero de Estudios: Julián Faivovich Buenos Aires, 2014 Fecha de defensa: 19 de diciembre de 2014 Comunicación química en anuros: una perspectiva integral a partir de aspectos comportamentales, morfológicos y químicos en dos especies de Hypsiboas (Amphibia: Anura: Hylidae) RESUMEN La piel de los anfibios es un órgano dinámico con una enorme diversidad de funciones. Una característica compartida por prácticamente todas las especies de anfibios es la presencia de dos tipos de glándulas dérmicas, mucosas y serosas. A pesar de la similitud estructural, estas glándulas pueden presentar una gran variabilidad morfológica alcanzando un alto grado de especialización en el cumplimiento de dos funciones, la comunicación química y la defensa contra predadores. Desde un aspecto bioquímico, el análisis de las secreciones glandulares posee un potencial enorme dado la diversidad de compuestos. Los estudios desarrollados en algunos de ellos (i.e., péptidos, alcaloides, aminas biogénicas) permite tener una idea aproximada de la variación estructural y funcional. Sin embargo, es limitado el conocimiento que se tiene sobre la química de los compuestos volátiles y de los componentes secretados por las glándulas de la piel sexualmente dimórficas (SDSGs); fuente potencial de feromonas. Tradicionalmente los estudios de comunicación en los anuros se focalizaron en el análisis de las señales acústicas, aunque existían reportes aislados de la participación de otras señales (i.e., visual, química, táctil) en algunas especies. Los trabajos recientes muestran que estas señales tienen una distribución taxonómica mucho mayor a la considerada inicialmente, en tanto que la co-ocurrencia de distintas señales sugieren la existencia de comunicación multimodal en algunas especies. En la presente tesis se abordaron aspectos morfológicos, químicos y comportamentales de las secreciones glandulares en dos especies de Hypsiboas, considerando un contexto reproductivo en H. punctatus y un contexto defensivo en H. pulchellus. Se analizó la estructura y la ultraestructura de la piel de las regiones mental, lateral, dorsal y ventral en machos y hembras de H. punctatus. Se identificaron dos tipos de SDSGs, en las regiones mental y lateral de los machos, representando el primer caso de co-ocurrencia de diferentes SDSGs en una misma especie de anuro. A partir de estos resultados se caracterizó la variación estructural e histoquímica, y se estudió la evolución de las glándulas mentales y laterales en 28 especies de Cophomantini, incluyendo representantes de prácticamente todos los grupos de especies. La ocurrencia de estas glándulas, al igual que la variación morfológica, presenta una distribución taxonómica compleja con diferentes niveles de homoplasia. A partir del análisis de las secreciones de las glándulas dorsales de machos y hembras se determinó la estructura primaria de 13 péptidos y se identificaron dos compuestos relaciones con alcaloides. Se identificó un péptido presente exclusivamente en las secreciones de las glándulas mental y lateral de los machos. Los estudios comportamentales en el campo y en condiciones controladas de laboratorio, mostraron que los machos de H. punctatus son territoriales y poseen un repertorio vocal complejo con al menos siete cantos diferentes asociados a contextos particulares. Las hembras elegirían a los machos en base a la información de las señales acústicas y químicas, lo que sugiere la existencia de comunicación multimodal en esta especie. El estudio morfológico e histoquímico de las glándulas de H. pulchellus evidenció la presencia de las dos glándulas comunes de los anfibios (mucosas y serosas). Sin embargo, los análisis de estructura, ultraestructura y de los mecanismos fisiológicos de descarga permiten inferir que las glándulas serosas cumplirían diversas funciones, entre ellas almacenar los componentes volátiles secretados en situaciones de estrés. Estas glándulas presentan una mayor concentración en la región escapular, lo que podría relacionarse con estrategias defensivas. A partir del análisis de las secreciones de las glándulas dorsales de machos y hembras se determinó la estructura primaria de 17 péptidos y se identificaron dos compuestos relaciones con alcaloides. Las secuencias peptídicas de H. pulchellus e H. punctatus presentan una gran variabilidad. Estos compuestos no fueron publicados anteriormente, aunque algunos de los péptidos presentaron una gran similitud con los descriptos en otras especies de Cophomantini. Se compararon las secuencias de todos los péptidos identificados hasta el momento en esta tribu. El solapamiento de las secuencias de los péptidos de H. pulchellus e H. punctatus con partes de péptidos mayores de otras especies sugiere la existencia de endopeptidasas específicas. Se caracterizó la fracción volátil de las secreciones de la región dorsal y lateral en H. pulchellus e H. riojanus. Se identificaron 42 compuestos (40 en H. pulchellus y 37 en H. riojanus) pertenecientes a nueve grupos funcionales, lo que constituye la mayor diversidad de compuestos volátiles identificados en una especie de anfibio. Los diferentes componentes tendrían distinto origen y función. Palabras claves: anuros, compuestos volátiles, comunicación multimodal, diversidad química, glándulas, péptidos. Chemical communication in anurans: an integral perspective from behavioral, morphological, and chemical aspects in two species of Hypsiboas (Amphibia: Anura: Hylidae) ABSTRACT The skin of amphibians is a dynamic organ with diverse functions. Almost all the species of this group of vertebrates possess two types of dermal glands, mucous and serous glands. Despite the structural similarities within each glandular type, they may have a great morphological variation getting a high degree of specialization related to chemical communication and defense against predators. From a biochemical perspective, and given the high diversity of their compounds, researches on the glandular secretions have an enormous potential. In particular, the studies performed on some of them (i.e., peptides, alkaloids, biogenic amines) allowed a general understanding of their structural and functional diversity. However, knowledge of the chemical composition of the volatile secretions, as well as from the secretion of sexually dimorphic skin glands (SDSGs; a likely source of sexual pheromones) in anurans, is very limited. Although there are isolated reports of the existence of different signals used for communication in some species (i.e., visual, chemical, tactile), studies of communication in anurans have traditionally focused in the analysis of acoustic signals. Nonetheless, recent investigations show that those considered rare signals have a wide taxonomic distribution, whereas the simultaneous occurrence of some of them suggest the existence of multimodal communication in several species. In this thesis are examined morphological, chemical and behavioral aspects from de glandular secretions of two species of Hypsiboas, considering a reproductive context in H. punctatus, and a defensive context in H. pulchellus. The structure and ultrastructure of skin from mental, lateral, dorsal, and ventral regions, was examined in males and females of H. punctatus. Two types of SDSGs were identified en mental and lateral regions of males. These results represent the first case of co-occurrence of different SDSGs in a single species. The structural and histochemical variation, as well as the evolution, of mental and lateral glands was analyzed in Cophomantini from samples of
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