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La Amígdala: Implicaciones Funcionales: M. Torras, I. Portell, I. Morgado
La Amígdala: Implicaciones Funcionales: M. Torras, I. Portell, I. Morgado
REVISIN
OBJETIVO
La amgdala esun grupo heterogneo de ncleos en cuanto a citoarquitectura, quimioarquitectura y conexiones, situados en el lbulo temporal de los mamferos [1-3]. Desde un punto d e vista funcional, se
considera que es una estructura clave en el sistema emocional del
cerebro [4]. Numerosos datos experimentales parecen indicar que la
amgdala se relaciona con los aprendizajes emocionales y con en el
almacenamiento de memorias afectivas [4,5]. Por otro lado, tambin
se ha sugerido que la amgdala podra ser un componentecrticodeun
sistemamoduladordelamemoria[6].Enesteartculo analizaremos los
datos que relacionan la amgdala con estas funciones, identificando
losdiferentessubsistemasdelamismaimplicadosencadaunade ellas.
471
DESARROLLO
La amgdala y los procesos emocionales
En los primeros experimentos en los que se realizaron lesiones del
lbulo temporal en primates, Klver y Bucy [7] constataron que se
produca un conjunto complejo de alteraciones en el comportamiento
social y emocional de los animales: despus de la lesin, los monos
considerados salvajes y fieros se mostraban dciles y no presentaban
las reacciones generalmente asociadas al miedo y a la ira. Se acercaban a hombres y animales que anteriormente les producan miedo y
tenan una tendencia generalizada a examinar los objetos con la boca
ms que con las manos, como si fueran incapaces de reconocer visualmente los objetos. Junto con estas alteraciones, tambin presentaban
hipersexualidad, a menudo dirigida indiscriminadamente, y dficit de
memoria. Posteriormente, en los aos 50, se demostr que los cambios en la conducta emocional observados en el sndrome KlverBucy se deban a la lesin de la amgdala [8].
Recibido: 06.04.01. Aceptado tras revisin externa sin modificaciones:06.05.01.
Departamento de Psicobiologa y Metodologa de las Ciencias de la Salud.
Facultad de Psicologa. Universidad Autnoma de Barcelona. Bellaterra,
Barcelona, Espaa.
Correspondencia: Dr. Ignacio Morgado Bernal. Departament de Psicobiologia
i Metodologia de les Cincies de la Salut. Facultat de Psicologia. Universitat
Autnoma de Barcelona. Apdo. de Correos 46. E-08193 Bellaterra, Barcelona.
M. TORRAS, ET AL
472
LA AMGDALA
473
M. TORRAS, ET AL
laadministracinsistmicadeadrenalina[72,73,101],nopareceafectar
a la adquisicin y retencin de ratas control entrenadas en tareas de
evitacin, en las que intervienen descargas elctricas y, por tanto, con
liberacin endgena de adrenalina. Es posible que, para activar la
estra terminal, sea necesario un determinado patrn de liberacin de
adrenalina, slo conseguido con la administracin exgena de adrenalina postentrenamiento o con el entrenamiento en algn tipo de
tarea o parmetros experimentales no utilizados en los estudios realizados hasta el momento [101]. Cabe destacar los resultados de un
experimento reciente realizado en nuestro laboratorio [101]: en las
ratas con lesin en la estra terminal que mostraron un bajo nivel de
aprendizaje en la sesin de adquisicin, el tratamiento postentrenamiento con adrenalina perifrica no slo bloque los efectos moduladores de esta hormona, sino que deterior la retencin a lo largo de
toda la sesin. Estos resultados podran explicarse si la adrenalina
reforzara los aspectos cognitivos de la tarea de aprendizaje aumentara
el estado emocional de los animales. As, el deterioro observado en
la prueba de retencin podra deberse a una potenciacin del miedo,
ya de por s elevado en estas ratas por el hecho de recibir una gran
cantidad de choques, sin ir acompaada de una facilitacin, a travs
de la estra terminal, de los aspectos cognitivos de la tarea.
Por ltimo, otra de las caractersticas que tendra que cumplir
la amgdala para poder considerarse un sistema modulador de la
memoria es la de modular el almacenamiento de la memoria en
otras zonas del encfalo, diferencindose de otros sistemas cerebrales que participan en la adquisicin y retencin de tareas relacionadas con un tipo especfico de memoria. En este sentido, los
resultados de algunos experimentos han sugerido que la amgdala
podra modular el almacenamiento de memorias dependientes del
hipocampo y del ncleo caudado [69,78,103-105].
CONCLUSIONES
Los resultados de diferentes experimentos apoyan la relacin de la
amgdala con los procesos emocionales. En este sentido, la amgdala
parece participar en la formacin de asociaciones entre estmulos y
reforzadores y, por tanto, en el proceso por el cual estmulos inicial-
menteneutrosadquierensignificacinemocionalymotivacionalcomo
resultado de la experiencia. En cambio, no est claro que la amgdala
sea el lugar de almacenamiento de este tipo de memorias o slo sea
necesaria para su expresin. En este sentido, algunos investigadores
han propuesto que la amgdala no sera el lugar de almacenamiento
de memorias, sino que formara parte de un sistema endgeno modulador de la memoria, que afectara a memorias almacenadas en diferentes regiones del cerebro. A nuestro entender, esta funcin de la
amgdala propuesta por McGaugh et al [6] es compatible con la
funcin de asignar contenido emocional a los estmulos que se asocian a una experiencia aversiva. Ambas funciones parecen estar bastante corroboradas por los datos experimentales y, de hecho, no nos
debera extraar que la misma estructura que se encarga de asignar
valor emocional a estmulos o situaciones inicialmente neutras sea
capaz de modular la memoria de los estmulos, que al haber adquirido
contenido emocional son emocionalmente relevantes. Adems, los
datos existentes sugieren que no sera todo el complejo amigdaloide
el implicado en ambas funciones, sino que los ncleos lateral y central
estaran implicados en la asociacin entre estmulos y reforzadores y
en la expresin de la respuesta emocional, respectivamente, mientras
que el ncleo basolateral estara sobre todo implicado en la modulacin de la memoria cuando se produce activacin emocional.
Aunque parece confirmado que la amgdala tiene una funcin
en la modulacin de la memoria, esto no es incompatible con el
hecho de que esta estructura sea el lugar de almacenamiento de las
memorias emocionales. La mayora de experimentos realizados
con el procedimiento de miedo condicionado sugieren que la amgdala es el lugar de almacenamiento de este tipo de memoria, mientras que los experimentos con aprendizajes de evitacin parecen
indicar que la amgdala slo es necesaria para la expresin de este
tipo de memorias. Recordemos que el procedimiento de evitacin
implica tanto un componente de condicionamiento clsico como el
aprendizaje de una respuesta instrumental, que podra almacenarse
en otras estructuras cerebrales diferentes a la amgdala. En este
sentido, podra ser que el aprendizaje emocional que tiene lugar en
la amgdala fuera el que modulara la memoria de tareas ms complejas almacenadas en otras regiones del encfalo.
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