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Redalyc.Understanding the Neurobiological Mechanisms of Learning and Memory: Memory Systems of the Brain, Long Term Potentiation

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Redalyc.Understanding the Neurobiological Mechanisms of Learning and Memory: Memory Systems of the Brain, Long Term Potentiation Salud Mental ISSN: 0185-3325 [email protected] Instituto Nacional de Psiquiatría Ramón de la Fuente Muñiz México Leff, Philippe; Romo, Héctor; Matus, Maura; Hernández, Adriana; Calva, Juan Carlos; Acevedo, Rodolfo; Torner, Carlos; Gutiérrez, Rafael; Anton, Benito Understanding the neurobiological mechanisms of learning and memory: Memory systems of the brain, long term potentiation and synaptic... Salud Mental, vol. 25, núm. 4, agosto, 2002, pp. 78-94 Instituto Nacional de Psiquiatría Ramón de la Fuente Muñiz Distrito Federal, México Available in: http://www.redalyc.org/articulo.oa?id=58242508 How to cite Complete issue Scientific Information System More information about this article Network of Scientific Journals from Latin America, the Caribbean, Spain and Portugal Journal's homepage in redalyc.org Non-profit academic project, developed under the open access initiative UNDERSTANDING THE NEUROBIOLOGICAL MECHANISMS OF LEARNING AND MEMORY: MEMORY SYSTEMS OF THE BRAIN, LONG TERM POTENTIATION AND SYNAPTIC PLASTICITY PART III B Philippe Leff1, Héctor Romo2, Maura Matus1, Adriana Hernández1, Juan Carlos Calva1, Rodolfo Acevedo1, Carlos Torner3, Rafael Gutiérrez2, Benito Anton1. SUMMARY RESUMEN One of the central issues in neuroscience is concerned with the El fenómeno de LTP es una forma de plasticidad sináptica activity-dependent synaptic plasticity in learning and memory. ampliamente aceptado como un modelo de estabilización de In such context, changing the strength of synaptic activity sinapsis en procesos neurobiológicos como el desarrollo del between neurons has been widely accepted as the mechanism SNC y el fenómeno de aprendizaje y memoria. Desde su des- responsible by which memory traces are encoded and stored cubrimiento por Bliss y Lomo (1973), el fenómeno de in the brain. Thus, the synaptic plasticity and memory hypothesis potenciación a largo plazo (PLP) o LTP (Long-Term Potentiation, (SPM hypothesis) shows that activity-dependent synaptic plasticity por sus siglas en inglés) ha sido definido convencionalmente is induced at appropiate synapses during memory formation, como la estimulación aferente de alta frecuencia que es capaz which is essential for information storage for the type of memory de despolarizar la célula postsináptica, a través de la activación involved in the brain area where plasticity is detected or observed. de receptores glutamaérgicos, con la resultante entrada de cal- Several criteria and experimental strategies are outlined, and cio a la neurona postsináptica. Este evento neurobiológico pro- used to investigate this hypothesis. Long-term potentiation (LTP) duce un incremento intracelular en la concentración de calcio as an experimental model to study the cellular basis of learning [(Ca)i] que induce la activación de diferentes sistemas moleculares and memory, is one of the most fascinating phenomena that de señalamiento intracelular (AMPc, proteínas cinasas, have raised a great interest in neuroscience. LTP is a form of fosforilación de proteínas intracelulares) que conlleva a una synaptic plasticity that is accepted as a cellular model for alteración de la actividad postsináptica y/o presináptica, dando stabilization of synapses in several neurobiological events such por resultado un persistente incremento de respuesta sináptica as development, learning and memory. Defined as the increase específica dependiente de la activación del receptor in the strength of synaptic transmission observed after tetanic glutamaérgico NMDA. Un alto porcentaje de los resultados stimulation, this phenomenon can be measured from hours to experimentales relativos al fenómeno de LTP se ha centrado days, and even outlast the stimulus that induced it over time. en las formas de LTP dependientes de la activación y función The role of LTP in learning has been a central issue in de este subtipo de receptor glutamaérgico, particularmente en neuroscience, particularly in studies focused on the NMDA re- la corteza cerebral, la formación hipocampal y las estructuras ceptor-dependent forms of LTP. Thus, much of the experimen- amigdalinas, estructuras neuroanatómicas que conforman el tal work performed in research regarding LTP has been aimed sistema límbico en los mamíferos. Sin embargo, han surgido to investigate if LTP equals memory. This review described several muchas interrogantes cuando se trata de igualar los eventos properties of synaptic plasticity as well as the neural substrates experimentales observados, del LTP, con los eventos de memo- where synaptic plasticity events are embedded in networks, so ria que ocurren en el cerebro de los mamíferos. Por ejemplo, as to establish the processing of learning and memory formation. de estas interrogantes podemos mencionar la relación que guardan las propiedades analizadas del LTP con respecto a la Key words: Long-term potentiation, long-term depression, función de la memoria, qué tipos de aprendizaje están relacio- synaptic plasticity, synaptic strength, memory, learning, nados con el desarrollo del fenómeno de LTP y que áreas hippocampus, amigdala. cerebrales se involucran en el desarrollo de este proceso. Si el 1 Laboratorio de Neurobiología Molecular y Neuroquímica de Adicciones. Instituto Nacional de Psiquiatría Ramón de la Fuente. Calzada México-Xochimilco 101. San Lorenzo Huipulco, 14370, México D.F. [email protected] 2 Departamento de Fisiología, Biofísica y Neurociencias. CINVESTAV. Apartado Postal 14-740, 07000. México D.F. [email protected] 3 Atención a la Salud, CBS, Universidad Autónoma Metropolitana-Xochimilco. Calzada del Hueso 1100, Col. Villa Quietud, 04960 México, D.F. 78 Salud Mental, Vol. 25, No. 4, agosto 2002 fenómeno del LTP juega un papel relevante en el desarrollo de dizaje, mediado a través de la facilitación o bloqueo de la la memoria, se debería postular, en principio, que la actividad- actividad sináptica o de la eficacia sináptica resultante. dependiente de la plasticidad sináptica y las diferentes formas Diversos estudios electrofisiológicos han demostrado que los de expresión de memoria que existen en el cerebro, compar- mecanismos neuronales involucrados, tanto en la inducción ten un denominador común. Es decir, que la actividad resul- del fenómeno de LTP como del fenómeno del LTD, en dife- tante de la plasticidad sináptica es inducida en sinapsis apro- rentes regiones del hipocampo, pueden ser dependientes o piadas durante la formación de cualquier evento o fenómeno independientes del receptor glutamaérgico, NMDA; pero am- de memoria analizado. Este proceso neurobiológico debe ser bos eventos implican la relación de la actividad presináptica relevante y suficiente para almacenar la información pertinente con una despolarización o hiperpolarización de la neurona al tipo de memoria mediada por una región cerebral específica, postsináptica. Más aún, dependiendo del grado de estimulación en la que ocurre un evento de plasticidad sináptica. La plastici- de los circuitos neuronales, responsables de inducir cambios dad sináptica es un evento neurofisiológico que induce patro- en la actividad sináptica o incrementos de la eficacia sináptica, nes específicos en la actividad neuronal, mediado por eventos en intervalos de tiempo definidos, en las sinapsis de las neuronas neuroquímicos y mecanismos moleculares que, finalmente, con- operantes, pueden detectarse cambios en la respuesta en la llevan a la generación de cambios en la excitabilidad neuronal actividad sináptica. Dichos cambios ocurren ocasionalmente y en la eficacia sináptica y que permanecen por muy largo (en una sola ocasión), con posterioridad a los procesos de tiempo y perduran indefinidamente en relación con los even- estimulación por el contacto entre neuronas presinápticas y tos neurobiológicos que los suscitan. En este contexto, se pue- postsinápticas. Estos resultados han permitido postular, emplean- de resumir que, tomando como base las propiedades de la do modelos de circuitos neuronales funcionales, la hipótesis plasticidad sináptica, el fenómeno de LTP, el fenómeno de sobre la existencia de “sinapsis silentes o silenciosas”. Esta hi- DAP (depresión a largo plazo) o LTD (Long-Term Depression, pótesis explica la transformación de sinapsis inactivas en sinapsis por sus siglas en inglés) constituyen los modelos fisiológicos activas mediante la síntesis e inserción de diferentes subtipos más viables y apropiados en la generación de diferentes siste- de receptores glutamaérgicos, por ejemplo, el subtipo de recep- mas de generación de memoria, tales como la codificación y tor AMPA que permite sustentar la vieja teoría sobre la expre- almacenamiento de información y la consolidación de trazos sión del fenómeno de LTP. Más aún, estudios recientes han de memoria perdurables en el tiempo. Diversos estudios expe- demostrado que la persistencia del fenómeno del LTP, en siste- rimentales han demostrado que el procesamiento de memoria mas neuronales en el SNC de mamíferos, es producto tanto de establecido por los mecanismos neurobiológicos que inducen la continua activación del receptor glutamaérgico, NMDA, como y mantienen el fenómeno de LTP o LTD, ocurre a través de la de la síntesis de novo de proteínas intracelulares esenciales que activación de circuitos neuronales específicos. En tales circui- consolidan los eventos de plasticidad sináptica dependientes tos, la codificación y almacenamiento de información (trazos de LTP. En el contexto de la plasticidad sináptica relacionada de memoria) ocurre como producto de las propiedades de los con los eventos biológicos de memoria y aprendizaje,
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