Learning and Memory: The Behavioral and Biological Substrates

By Isidore Gormezano; Edward A. Wasserman | Go to book overview

16
The Essential Memory Trace Circuit for a Basic Form of Associative Learning

Richard F. Thompson University of Southern California

Joseph E. Steinmetz Indiana University

Some years ago we selected classical conditioning of the nictitating membrane/eyelid closure response as a model system in which to analyze the neuronal substrates of basic associative learning and memory phenomena. We adopted this paradigm and the rabbits as the experimental animal of choice for two key reasons: (a) There is an extensive literature on the properties and parameters of this form of associative learning in both humans and animals (particularly the rabbit) ( Black and Prokasy, 1972; Gormezano, 1972); and (b) it obeys the basic "laws" and exhibits the fundamental phenomena of associative learning in a similar manner in humans and in other mammals.


THE CEREBELLUM AND MOTOR LEARNING

When we began this work about 18 years ago, we had no idea that we would be led to the cerebellum as the key structure that appears to store the essential memory trace. With the advantage of hindsight, it is perhaps not so surprising. The conditioned eyelid closure response is a very precisely timed movement-- over CS--US onset intervals ranging from about 100 msec to over 1 sec, the learned response develops such that the eyelid closure is maximal at the time of onset of the US. In this sense it is a maximally adaptive response. (By "adaptive" we refer only to the temporal properties of the CR relative to the onset of the US and not to issues relating to reinforcement or the "law of effect"; see Gormezano & Coleman, 1973). It is also a very precisely timed "skilled" movement, perhaps the most elementary form of learned skilled movement. Our results strongly support the general spirit of earlier theories of the role of the

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