is repeated again and again.

This modularity raises the possibility that,

although different regions of the cerebellum receive different input, each
region might perform a similar set of operations on that information before
passing the results back out to the rest of the nervous system (Boyden,
Katoh, & Raymond, 2004).

Eyeblink Classical Conditioning as a Prototype of Cerebellum-

Dependent Memory

The cerebellum is important for a range of sensorimotor functions,

including adaptations of the vestibulo-occular response and coordinated
motor skill learning. The cerebellum has also been associated with
nonmotor learning and other examples of cognition and behavior. The
present section focuses on the best-understood example of cerebellar-
dependent learning, eyeblink classical conditioning, as a framework for
highlighting the role of the cerebellum in memory based on the idea that the
modular circuitry of the cerebellum offers a generalized capacity for
processing of specific inputs.
In standard “delay” eyeblink classical conditioning, a conditioned
stimulus (CS; e.g., a brief tone) is initiated immediately (typically a second
or less) prior to the delivery of an unconditioned stimulus (US; e.g., a brief
puff of air to the eye), and both stimuli then terminate at the same time. The
US triggers a reflexive, unconditioned eyeblink (the UR), and after many
pairings of CS and US, the CS eventually elicits an eyeblink as a
conditioned response timed such that the blink is most robust at the US
onset. Remarkably, rabbits with no cerebral cortex, basal ganglia, limbic
system, thalamus, or hypothalamus showed normal retention of the
conditioned eyeblink response (Mauk & Thompson, 1987), indicating that
no forebrain structures were an essential part of the neural circuitry
supporting standard eyeblink conditioning.
Numerous studies by Thompson and colleagues have traced the paths
to the cerebellum taken by incoming sensory information about the CS and
the US and have found that these paths converge on one particular
cerebellar nucleus, the interpositus nucleus, and that outputs of this nucleus
have a clear route to producing conditioned eyeblink responses (Fig. 48.13;
Thompson, 2005). These experiments were able to capitalize on the well-