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Centros Catecolaminérgicos En El Sistema Nervioso Central De Anfibios: Estudio Hodológico Y Neuroquímico

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Centros Catecolaminérgicos En El Sistema Nervioso Central De Anfibios: Estudio Hodológico Y Neuroquímico UNIVERSIDAD COMPLUTENSE DE MADRID FACULTAD DE CIENCIAS BIOLÓGICAS Departamento de Biología Celular CENTRO CATECOLAMINÉRGICOS EN EL SISTEMA NERVIOSO CENTRAL DE ANFIBIOS: ESTUDIO HODOLÓGICO Y NEUROQUÍMICO MEMORIA PARA OPTAR AL GRADO DE DOCTOR PRESENTADA POR Cristina Sánchez – Camacho Blázquez Bajo la dirección del doctor Agustín González Gallegos Madrid, 2002 ISBN: 84-669-1718-7 Universidad Complutense de Madrid Facultad de Ciencias Biológicas Centros Catecolaminérgicos en el Sistema Nervioso Central de Anfibios. Estudio hodológico y neuroquímico Cristina Sánchez-Camacho Blázquez 2002 Universidad Complutense de Madrid Facultad de Ciencias Biológicas Centros Catecolaminérgicos en el Sistema Nervioso Central de Anfibios. Estudio hodológico y neuroquímico Trabajo de Investigación que presenta Cristina Sánchez-Camacho Blázquez para optar al grado de Doctor en Ciencias Biológicas en la Universidad Complutense de Madrid Fdo. Dña. Cristina Sánchez-Camacho Blázquez Dirigido por el Doctor Agustín González Gallegos Profesor Titular del Departamento de Biología Celular de la Facultad de Ciencias Biológicas de la Universidad Complutense de Madrid Fdo. D. Agustín González Gallegos D. Agustín González Gallegos, Profesor Titular del Departamento de Biología Celular de la Facultad de Ciencias Biológicas de la Universidad Complutense de Madrid, HACE CONSTAR: Que Doña Cristina Sánchez-Camacho Blázquez ha realizado bajo mi dirección el trabajo de su Tesis Doctoral “Centros catecolaminérgicos en el Sistema Nervioso Central de anfibios. Estudio hodológico e inmunohistoquímico”, que ha terminado con el mayor aprovechamiento. Revisado el presente trabajo, quedo conforme con su presentación para ser juzgado. Y para que conste y surta los efectos oportunos, lo firmo en Madrid a dieciocho de septiembre de dos mil dos. Fdo. D. Agustín González Gallegos A mi padre A mi madre A Jorge Índice Índice 1 Introducción General 13 2 Conexiones aferentes de la médula espinal 27 Descending supraspinal pathways in amphibians. I. A dextran amine tracing study of their cells of origin 29 3 Inervación catecolaminérgica de la médula espinal 51 Descending supraspinal pathways in amphibians. II. Distribution and origin of the catecholaminergic innervation of the spinal cord 53 4 Desarrollo de las conexiones aferentes y catecolaminérgicas de la médula espinal 75 Descending supraspinal pathways in amphibians. III. Development of descending projections to the spinal cord in Xenopus laevis with emphasis on the catecholaminergic inputs 77 Origin and development of descending catecholaminergic pathways to the spinal cord in amphibians 91 5 Inervación catecolaminérgica del techo óptico 99 Distribution and origin of the catecholaminergic innervation in the amphibian mesencephalic tectum 101 6 Inervación catecolaminérgica de la región septal 115 Catecholaminergic innervation of the septum in the frog: a combined immunohistochemical and tract-tracing study 117 7 Resumen de los resultados y Discusión general 131 8 Conclusiones 151 9 Agradecimientos 155 Capítulo 1 Introducción General Síntesis y Metabolismo de las Catecolaminas Técnicas de Detección de las Catecolaminas Organización de los Sistemas CA en el SNC de Mamíferos Organización de los Sistemas CA en el SNC de Anfibios El Modelo Neuromérico en el Estudio de la Distribución de los Grupos CA Aspectos Filogenéticos de los Sistemas CA en el SNC de Vertebrados Objetivos y Metodología CAPÍTULO 1. INTRODUCCIÓN GENERAL 15 Desde la demostración de su presencia en el sistema ner- vioso central (SNC) en los años cincuenta, las catecolaminas han recibido gran atención, particularmente debido a su impli- cación en determinados procesos patológicos y desórdenes neurológicos como la enfermedad de Parkinson, estados de ansiedad y depresión o la esquizofrenia. Asimismo, se han acumulado un gran número de datos acerca de la organización de los sistemas catecolaminérgicos (CA) en el SNC de los vertebrados amniotas (mamíferos, aves y reptiles) y anamnio- tas (anfibios y peces). La distribución de este grupo de neuro- transmisores parece altamente conservada a lo largo de la evo- lución y constituye posiblemente uno de los sistemas neuro- químicos filogenéticamente más antiguos del encéfalo de ver- tebrados (Parent, 1984). Sin embargo, existen pocos datos acerca de la conectividad de sus grupos celulares que permitan establecer homologías claras no sólo en función de su topogra- fía sino también en base a su hodología y ontogenia. Síntesis y Metabolismo de las Catecolaminas Las catecolaminas (CA) son un grupo de compuestos or- gánicos formados por un anillo de benzeno con dos grupos hidroxilo adyacentes (grupo catecol) y una cadena amina en el lado opuesto. Las principales CA usadas por el sistema nervio- so son la dopamina (DA), la noradrenalina (NA) y la adrenali- na (A), derivadas del aminoácido L-tirosina. La primera enzi- ma en la ruta de síntesis de las catecolaminas es la tirosina hidroxilasa (TH) que convierte la L-tirosina derivada de la dieta en la primera catecolamina de la ruta, la L-DOPA (L- dihidroxifenilalanina), tras la adición de un grupo hidroxilo al Fig. 1. Ruta de síntesis de las catecolaminas. anillo catecol (Fig. 1). Ésta es la enzima limitante de la reac- ción y está presente en todas las neuronas catecolaminérgicas. A continuación, la DOPA es transformada en DA mediante la recaptación de la DA (DAT). Este proceso permite la reutiliza- descarboxilación del grupo amino en una reacción catalizada ción de las CA evitando su degradación extraneuronal, al mis- por la enzima DOPA descarboxilasa (también conocida como mo tiempo que controla su concentración en el espacio extra- L-aminoácido aromático descarboxilasa, AACD). En las neu- celular regulando el nivel de activación de los receptores post- ronas noradrenérgicas, la DA actúa como precursor en la sínte- sinápticos. sis de la NA (o norepinefrina) tras la adición de un grupo Finalmente, en el proceso de inactivación de las CA parti- hidroxilo a la cadena lateral por la enzima dopamina β- cipan varias enzimas catabólicas, entre las que se incluyen la hidroxilasa (DBH). Finalmente, la NA es utilizada como pre- monoamino oxidasa (con dos isoformas, la MAO-A y MAO- cursor del producto final de la ruta metabólica de las catecola- B) y la catecol-O-metiltransfersa (COMT). La MAO se en- minas, la adrenalina (o epinefrina). Esta última es sintetizada cuentra en neuronas y células gliales y debido a su localización por la enzima feniletanolamina N-metiltransferasa (PNMT), intracelular, juega un papel preponderante en la degradación de tras la metilación del grupo amino de la cadena lateral de la las CA que no están almacenadas en vesículas. La COMT está NA (Reiner, 1994). presente virtualmente en casi todas las células unida a la mem- En las células dopaminérgicas, la conversión de la tirosina brana postsináptica, e inactiva las CA que puedan escapar de la en L-DOPA y de ésta en DA ocurre en el citoplasma neuronal. recaptación por los terminales sinápticos (Reiner, 1994). Por el contrario, en las neuronas noradrenérgicas la DA pasa a las vesículas de almacenamiento donde sufre la β- Técnicas de Detección de las Catecolaminas hidroxilación para dar lugar a la NA. Así, la mayor parte de la Las técnicas de histofluorescencia desarrolladas a finales enzima DBH está unida a la membrana vesicular en los termi- de la década de los sesenta, fueron pioneras en el estudio de la nales nerviosos. Finalmente, la NA es transformada en A en el distribución de las catecolaminas en el SNC. En particular, el citoplasma de las neuronas adrenérgicas que contienen la en- método de fluorescencia inducida por formaldehído, desarro- zima PNMT. llado por Falck y cols. (1962), y sus posteriores modificaciones Tras su liberación, las CA actúan sobre sus receptores de están basadas en las características químicas de las aminas que membrana localizados en las neuronas postsinápticas. Existen experimentan con facilidad reacciones de oxido-reducción y dos tipos de receptores para DA, D1 (con los subtipos D1 y condensación. Así, se observó que las catecolaminas y la sero- D5) y D2 (D2, D3 y D4), mientras que los efectos de la NA y tonina daban lugar a productos fluorescentes de color verde o la A están mediados a través de los receptores α1-adrenérgicos amarillo respectivamente, en presencia de formaldehído. Sin (α1A, α1B, α1D), α2-adrenérgicos (α2A, α2B, α2C) o β- embargo, esta metodología presenta varias limitaciones con adrenérgicos (β1, β2, β3). Por otro lado, tras la liberación de respecto a la sensibilidad y especifidad del marcaje de las CA. las CA a la hendidura sináptica se produce un proceso de re- Así, el principal inconveniente de estas técnicas es que no captación dependiente de energía mediante un transportador permiten distinguir entre las diferentes catecolaminas presen- localizado en la membrana externa del terminal. Existe un tes, además de la inestabilidad de los productos formados y su transportador específico para la NA (NET) que se encuentra baja sensibilidad. sólo en las neuronas noradrenérgicas, y otro para la 16 CAPÍTULO 1. INTRODUCCIÓN GENERAL Con el desarrollo de las técnicas de inmunohistoquímica en rescencia. En este trabajo se describen 12 grupos CA diferen- los años ochenta, mediante la utilización de anticuerpos contra tes que denominaron de A1 a A12 de caudal a rostral. Estudios las catecolaminas o sus enzimas de síntesis, se obtuvo una posteriores empleando técnicas inmunohistoquímicas, han herramienta más sensible y específica para su localización en permitido determinar qué
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