Invertebrate Studies: Key Insights from

Aplysia into Basic Mechanisms of

Learning

Since the mid-1960s, the marine mollusc Aplysia has proven to be an

extremely useful model system to gain insights into the neural and
molecular mechanisms of simple forms of memory. Indeed, the pioneering
discoveries of Eric Kandel using this animal were recognized by his receipt
of the Nobel Prize in Physiology or Medicine in 2000. A number of
characteristics make Aplysia well suited for the examination of the
molecular, cellular, morphological, and network mechanisms underlying
neuronal modifications (plasticity) and learning and memory. The animal
has a relatively simple nervous system with large, individually identifiable
neurons that are accessible for detailed analyses. Indeed, the neurons and
neural circuits that mediate many behaviors in Aplysia have been identified.
In several cases, these behaviors have been shown to be modifiable by
learning. Moreover, specific loci within neural circuits at which
modifications occur during learning have been identified, and aspects of the
cellular mechanisms underlying these modifications have been analyzed
(For reviews see Bailey & Kandel, 2008; Byrne & Kandel, 1996;
Fioravante, Antzoulatos, & Byrne, 2008; Kandel, 2001).
Sensitization of withdrawal reflexes has been particularly well studied.
A single sensitizing stimulus, such as a brief several second-duration
electric shock, can produce a reflex enhancement that lasts minutes (short-
term sensitization), whereas prolonged training (e.g., multiple stimuli over
an hour or more) produces an enhancement that lasts from days to weeks
(long-term sensitization).

Multiple Cellular Processes Mediate Short- and

Long-Term Sensitization in Aplysia