Journal of Neuroscience, Vol 7, 120-132, Copyright © 1987 by Society for Neuroscience
Development of learning and memory in Aplysia. I. Functional assembly of gill and siphon withdrawal
CH Rankin, M Stopfer, EA Marcus and TJ Carew
The marine mollusc Aplysia californica provides an excellent preparation
with which to examine the development of the neuronal control of behavior
for 2 reasons: first, adult Aplysia exhibit a variety of behaviors that are
well understood in cellular terms; and second, the development of Aplysia
from embryo to adult has been studied in considerable detail. Among the
best understood behaviors in Aplysia are the withdrawal responses of the
mantle organs (the gill, siphon, and mantle shelf), which exhibit 2
different kinds of behaviors: "spontaneous" contractions that are part of a
fixed action pattern, a respiratory pumping sequence of the mantle organs,
and reflex contractions in response to tactile stimuli. We have examined
the development of both of these withdrawal behaviors in juvenile stages
9-12 and found that they are functionally assembled according to different
ontogenetic timetables. Spontaneous contractions. As soon as the siphon and
gill emerge, in stages 9 and 10, respectively, they each show a high rate
of spontaneous contraction that gradually diminishes throughout subsequent
stages until it reaches the low rate typical of adults (stage 13). Since
the siphon emerges first, it already exhibits a significant decline in its
spontaneous activity (e.g., in stage 11) when the gill's spontaneous
activity is at its highest. In addition to a developmental trend in the
rate of contractions, there was also a clear developmental progression in
the degree of cocontraction of the siphon and gill during spontaneous
contractions. In adults, the siphon and gill show a very high degree of
cocontraction during spontaneous pumping. However, in juvenile animals,
there was a very low degree. Thus, it appears that the siphon and gill
withdrawal components of the fixed action pattern become progressively more
functionally coupled during juvenile development. Reflex contractions. As
soon as the siphon and gill emerge in their respective developmental
stages, they exhibit a brisk withdrawal reflex to tactile stimulation of
the siphon. Moreover, at each developmental stage, reflex siphon
contractions were graded as a function of stimulus intensity, as they are
in the adult. Finally, throughout development tactile stimulation of the
siphon invariably evoked coincident contractions of both the siphon and the
gill, which is characteristic of the adult reflex. Thus, unlike the fixed
action pattern that takes several weeks to mature, the defensive withdrawal
reflex closely resembles the adult form as soon as the effector organs
emerge during juvenile development.(ABSTRACT TRUNCATED AT 400 WORDS)