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Estudio De La Morfología Y El Desarrollo Del Tracto De Salida Cardíaco En Xenopus Laevis

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Estudio De La Morfología Y El Desarrollo Del Tracto De Salida Cardíaco En Xenopus Laevis Facultad de Ciencias Departamento de Biología Animal Programa de doctorado: Biología Celular y Molecular TESIS DOCTORAL Estudio de la morfología y el desarrollo del tracto de salida cardíaco en Xenopus laevis. Implicaciones evolutivas Agustina María Torres Prioris 2017 Directora Ana Carmen Durán Boyero UNIVERSIDAD DE MÁLAGA FACULTAD DE CIENCIAS Departamento de Biología Animal TESIS DOCTORAL: Estudio de la morfología y el desarrollo del tracto de salida cardíaco en Xenopus laevis. Implicaciones evolutivas. Agustina María Torres Prioris Directora: Dra. Ana Carmen Durán Boyero Málaga, 2017 ! AUTOR: Agustina María Torres Prioris http://orcid.org/0000-0003-1532-1678 EDITA: Publicaciones y Divulgación Científica. Universidad de Málaga ! Esta obra está bajo una licencia de Creative Commons Reconocimiento-NoComercial- SinObraDerivada 4.0 Internacional: http://creativecommons.org/licenses/by-nc-nd/4.0/legalcode Cualquier parte de esta obra se puede reproducir sin autorización pero con el reconocimiento y atribución de los autores. No se puede hacer uso comercial de la obra y no se puede alterar, transformar o hacer obras derivadas. Esta Tesis Doctoral está depositada en el Repositorio Institucional de la Universidad de Málaga (RIUMA): riuma.uma.es A mis padres A mi abuelita A mis hermanos A Miguel “Yo no le canto a la luna porque alumbra nada más, le canto porque ella sabe Atahualpa Yupanqui de mi largo caminar” Índice ÍNDICE Prefacio ............................................................................................................... 1 Agradecimientos ................................................................................................. 3 Introducción ........................................................................................................ 9 Seno venoso ..................................................................................................... 13 Atrios ................................................................................................................. 15 Ventrículo .......................................................................................................... 17 El tracto de salida cardíaco ............................................................................... 18 Desarrollo cardíaco ........................................................................................... 24 La cardiogénesis y su regulación molecular ..................................................... 27 Objetivos del presente estudio .......................................................................... 31 Material y Métodos ............................................................................................. 33 Material ............................................................................................................. 35 Métodos ............................................................................................................ 36 Obtención del material ................................................................................ 36 Fecundación natural ............................................................................. 36 Fecundación in vitro ............................................................................. 37 Fijación de las muestras .............................................................................. 40 Técnicas histológicas e histoquímicas para microscopía óptica ................. 41 Hematoxilina de Delafield-eosina amarillenta ...................................... 44 Tricrómico de Mallory ........................................................................... 44 Tricrómico de Masson-Goldner ............................................................ 45 Resorcina-fucsina ................................................................................. 45 Ácido peryódico de Schiff (PAS) .......................................................... 46 Picrosirio con microscopía de polarización .......................................... 46 Azul alciano (Scott y Dorling, 1965) ..................................................... 47 Técnicas inmunohistoquímicas ................................................................... 48 Método de la inmunoperoxidasa .......................................................... 49 Método de la inmunofluorescencia ....................................................... 50 Método de inmunofluorescencia in toto ................................................ 52 Morfología de superficie mediante microscopía electrónica de barrido ...... 53 Cortes semifinos ......................................................................................... 54 Hibridación in situ ........................................................................................ 55 Reconstrucciones tridimensionales ............................................................. 58 Resultados ........................................................................................................... 61 Anatomía e histología del corazón adulto de Xenopus laevis ........................... 63 Componente proximal del tracto de salida ........................................................ 69 Componente distal del tracto de salida ............................................................. 88 Arterias coronarias en el tracto de salida .......................................................... 97 Desarrollo embrionario del corazón .................................................................. 100 Estadios 26-28 ............................................................................................. 101 Estadios 29-30 ............................................................................................. 104 Estadios 31-32 ............................................................................................. 105 Estadios 33-34 ............................................................................................. 109 Estadios 35-36 ............................................................................................. 109 Estadios 37-38 ............................................................................................. 115 Estadio 39 .................................................................................................... 118 Estadio 40 .................................................................................................... 122 Estadio 41 .................................................................................................... 125 Estadio 42 .................................................................................................... 128 Estadio 43 .................................................................................................... 129 Estadio 44 .................................................................................................... 134 Estadio 45 .................................................................................................... 135 Estadio 46 .................................................................................................... 140 Estadio 47 .................................................................................................... 144 Estadio 48 .................................................................................................... 146 Estadios 49-50 ............................................................................................. 150 Estadios 51-52 ............................................................................................. 152 Metamorfosis e individuos juveniles ............................................................ 156 Discusión ............................................................................................................. 161 Aspectos morfológicos del tracto de salida ....................................................... 169 Aspectos embriológicos .................................................................................... 181 Aspectos evolutivos ........................................................................................... 188 Pigmentación del corazón ................................................................................. 191 Conclusiones ...................................................................................................... 195 Conclusions ........................................................................................................ 201 Summary .............................................................................................................. 207 Bibliografía .......................................................................................................... 229 Prefacio Las investigaciones que han conducido a la redacción de la presente Tesis Doctoral se han realizado en el Departamento de Biología Animal de la Universidad de Málaga, en el ámbito de las actividades del grupo “Estudios Cardiovasculares en Vertebrados” - BIO 203 del Plan Andaluz de Investigación, Desarrollo e Innovación (PAIDI) de la Junta de Andalucía-. En el transcurso de este trabajo, la autora ha sido beneficiaria de una beca del Subprograma de Formación del Personal Investigador (Subdirección General de Formación y Movilidad de Investigadores, Dirección General de Investigación y Gestión del Plan Nacional de I+D+i), referencia BES-2011-04690, desde el 1 de septiembre de 2011 hasta el 31 de agosto de 2015, concedida por el Ministerio de Ciencia e Innovación. Además, ha recibido una Ayuda al Estudio del Personal Investigador en Formación del
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