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Las “Medusas” Como Depredadores Y Presas LAS “MEDUSAS” COMO DEPREDADORES Y PRESAS. ESTUDIO DE SU METABOLISMO RESPIRATORIO Y COMPOSICIÓN BIOQUÍMICA. JELLYFISH AS PREDATORS AND PREY: STUDYING THEIR RESPIRATORY METABOLISM AND BIOCHEMICAL COMPOSITION. Doctorado en Oceanografía y Cambio Global Daniel Rickue Bondyale Juez Universidad de Las Palmas de Gran Canaria Las Palmas de Gran Canaria Diciembre 2020 ii LAS “MEDUSAS” COMO DEPREDADORES Y PRESAS. ESTUDIO DE SU METABOLISMO RESPIRATORIO Y COMPOSICIÓN BIOQUÍMICA. “JELLYFISH” AS PREDATORS AND PREY: STUDYING THEIR RESPIRATORY METABOLISM AND BIOCHEMICAL COMPOSITION. Memoria presentada para optar al título de Doctor en Oceanografía y Cambio Global Doctoral dissertation for the Ph. D. degree in oceanography and Global Change Dirigida por los Doctores: May Gómez Cabrera y Theodore Train Packard Instituto de Oceanografía y Cambio Global (IOCAG) Instituto Universitario de Acuicultura Sostenible y Ecosistemas Marinos (ECOAQUA) – Grupo de Ecofisiología de organismos marinos (EOMAR) Facultad de Ciencias del Mar de la ULPGC Universidad de Las Palmas de Gran Canaria Las Palmas de Gran Canaria - Diciembre 2020 iii iv D. SANTIAGO HERNÁNDEZ LEÓN COORDINADOR DEL PROGRAMA DE DOCTORADO EN OCEANOGRAFÍA Y CAMBIO GLOBAL DE LA UNIVERSIDAD DE LAS PALMAS DE GRAN CANARIA, INFORMA, Que la Comisión Académica del Programa de Doctorado, en su sesión de fecha ....................... tomó el acuerdo de dar el consentimiento para su tramitación, a la tesis doctoral titulada " Las “medusas” como depredadores y presas. Estudio de su metabolismo respiratorio y composición bioquímica” presentada por el doctorando D. Daniel Rickue Bondyale Juez y dirigida por la Doctora May Gómez Cabrera y codirigida por el doctor Theodore T. Packard. Y para que así conste, y a efectos de lo previsto en el Artº 11 del Reglamento de Estudios de Doctorado (BOULPGC 7/10/2016) de la Universidad de Las Palmas de Gran Canaria, firmo la presente en Las Palmas de Gran Canaria, a...de.............................................de dos mil............ v vi UNIVERSIDAD DE LAS PALMAS DE GRAN CANARIA ESCUELA DE DOCTORADO Programa de doctorado en Oceanografía y Cambio Global Título de la Tesis LA“ MEDU“A“ COMO DEPREDADORE“ Y PRE“A“. E“TUDIO DE “U METABOLI“MO RESPIRATORIO Y COMPOSICIÓN BIOQUÍMICA. Tesis Doctoral presentada por D. Daniel Rickue Bondyale Juez Dirigida por el Dra. May Gómez Cabrera Codirigida por el Dr. Theodore T. Packard El/la Director/a, El/la Codirector/a El/la Doctorando/a, (firma) (firma) (firma) Las Palmas de Gran Canaria, a de de 20 vii viii A mis “mitocondrias”, ix AGRADECIMIENTOS/ACKNOWLEDGEMENTS Quiero agradecer la oportunidad al programa de ayudas para la formación de profesorado universitario del Ministerio de Educación y Formación Profesional, a la Universidad de Las Palmas de Gran Canaria, a la Facultad de Ciencias del Mar y al programa de doctorado de oceanografía y cambio global. Contar con programas e instituciones públicas destinados a que las personas tengamos acceso a la investigación y a la educación es un privilegio que valoro y querría que fuese perenne. La criptica dedicatoria, “A mis mitocondrias” viene motivada por la relación que tienen estos orgánulos con esta tesis y con la generación de energía. Pero, evidentemente, la dedicación es sobre todo figurada, no literal, y se refiere a aquellas personas que han sido mi fuente personal de energía. No de una energía que pueda ser medida o cuantificada, pero de una energía emocional sin la cual la autosuperación, en mi opinión, es imposible. En lugar de Adenosine triphosphate, el ATP del que me proveen estas mitocondrias emocionales sería Apoyo- Talento-Pasión, Atención-Tiempo-Paciencia, Amistad-Trabajo-Preguntas, Autoestima- Tenacidad-Perseverancia, Alegrías-Triunfo-Positividad... He intentado usar este ATP especial para convertirlo en este trabajo por lo que una parte de todas esas personas está aquí presente. Hay muchas “mitocondrias” incansables a las que agradecer. El trabajo aquí presentado pertenece sobre todo a la familia del grupo EOMAR. Quiero dar las gracias a May, Ted, Ico, Vane y Mayte, personas con las que he pasado muchas horas y cuya influencia y dedicación está presente en esta tesis. No quiero dejar fuera del agradecimiento también a todas las demás personas que forman o han formado parte de la familia EOMAR en algún momento ya que su compañía siempre ha sido agradable y estimulante. Agradezco también la labor de numerosos profesores a lo largo de mi vida cuyas lecciones individuales no recuerdo como no recuerdo todas las comidas que he comido pero de igual manera han contribuido seguro a lo que soy hoy. De igual, o tal vez, mayor medida me siento enormemente afortunado por el apoyo de amigos y familia cuyos nombres llenarían muchas páginas. I will also like to thank all the people which have provided me of opportunities to experience collaboration and integration in the scientific community. A special thanks goes to Jennifer Purcell whose contributions to the jellyfish research community have been extensive and it has been a privilege to collaborate with her in some publications. Also I acknowledge the participation of Javier Almunia and Ester Alonso of the Loro Parque Foundation who in several occasions have provided the organisms to do this research. Anna Belcher and Kathryn Cook have also been instrumental to experience a collaboration with other institutions I thank them and their colleagues for their positive and very inclusive attitudes, especially Nathan Hubbot a fellow “medusologist” who opened the doors of his house without hesitation before even knowing me. I will also like to mention all the people that received us in their institutions during our time in the United States and Mexico it was wonderful observing the work of other research centres and being able to share. Finally, I would like to thank Alexandra Elbakyan whose effort to make research more accessible has been vital. x “In the end respiration and burning are equivalent; the slight delay in the middle is what we know as life.” Nick Lane, The vital Question “The real world is in a much darker and deeper place than this, and most of it is occupied by jellyfish and things. We just happen to forget all that. Don't you agree? Two-thirds of earth's surface is ocean, and all we can see with the naked eye is the surface: the skin.” Haruki Murakami, The Wind-Up Bird Chronicle xi xii RESUMEN El estudio del zooplancton ha sido siempre bastante crustaceocéntrico lo cual no es de extrañar ya que son componentes mayoritarios del zooplancton y sus exosqueletos les permiten no ser dañados tan gravemente durante los muestreos. Sin embargo, esto ha contribuido a que otros grupos también abundantes pasen más desapercibidos como es el caso del plancton gelatinoso. Los organismos de este grupo con su alto contenido en agua tienen una gran fragilidad. Además, el plancton gelatinoso puede alcanzar tamaños grandes que limitan su captura por métodos tradicionales de muestreo de plancton y aquellos muestreados son fácilmente dañados o destruidos en el proceso. Junto a esto a la hora de estudiar el contenido digestivo de un hipotético depredador, un organismo gelatinoso se descompondrá más rápidamente que, por ejemplo, el exoesqueleto de otras presas del grupo de los crustáceos. Todo ello ha contribuido al menosprecio del plancton gelatinoso tanto en su rol como depredador planctónico como de presa. En este trabajo se busca contribuir al, a veces escaso, conocimiento sobre algunos miembros del plancton gelatinoso y su rol como depredador y presa. En concreto, para estimar cuantitativamente el consumo de presas necesario para estos organismos se estudió su metabolismo respiratorio. Durante el trabajo se revisaron métodos convencionales de medida de respiración a través de la tasa de consumo de oxígeno empleando incubaciones y monitorizando la concentración de O2 mediante el método Winkler, electrodos u optodos. Luego estos resultados se compararon con medidas de la respiración empleando técnicas enzimáticas que no requieren de incubación ni de muestreo del organismo vivo como es el análisis cinético del sistema de transporte de electrones que controlan la actividad respiratoria a nivel celular. Con ello se buscó validar e incorporar una metodología alternativa que permitiese trabajar con estos frágiles organismos aun estando dañados durante el muestreo. A través del estudio de la respiración pudimos cuantificar la cantidad de carbono necesaria para satisfacer esta actividad metabólica. Además, se emplearon estas medidas para no solo investigar el flujo y la transformación de carbono sino también para estudiar las conversiones a nivel energético que tienen lugar en el medio marino. Por otro lado, para entender su importancia como presa se analizó su contenido en sustancias orgánicas como proteínas, lípidos y carbohidratos. El contenido en dichas sustancias se empleó también para estimar su contenido energético y arrojar luz sobre la depredación cada vez más estudiada que sufren estos organismos con un mayor contenido en agua que otras presas. Específicamente se trabajó sobre todo con dos especies de escifozoos (Aurelia aurita y Pelagia noctiluca) y dos especies de hidrozoos (Physalia physalis y Velella velella). Estas últimas han sido rara vez estudiadas a nivel de fisiología respiratoria o contenido orgánico. Por tanto, en este trabajo seguramente están presentes las primeras medidas documentadas de respiración y composición en dichos integrantes del pleuston. Por otro lado, el estudio de la respiración y composición de estos organismos nos ha permitido documentar los cambios sufridos a nivel bioquímico y metabólico además de morfológico durante la transición de Aurelia aurita de larva a juvenil (éfira a medusa) y durante prolongados periodos de inanición. Estos estudios ecofisiológicos podrían explicar en parte algunos de los mecanismos que permiten la supervivencia en condiciones adversas, como por ejemplo la inanición. Seguramente estos procesos han contribuido a que este tipo de organismos haya logrado un importante éxito evolutivo al sobrevivir con un diseño muy parecido desde los comienzos de la vida eucariota pluricelular compleja siendo uno de los primeros metazoos.
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