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The Journal of Neurophysiology Vol. 81 No. 2 February 1999, pp. 654-667
Copyright ©1999 by the American Physiological Society
Department of Molecular and Integrative Physiology and the Neuroscience Program, University of Illinois, Urbana, Illinois 61801
Central pattern generator for escape swimming in the notaspid sea slug
Pleurobranchaea californica. Escape swimming in the notaspid opisthobranch Pleurobranchaea is an episode of
alternating dorsal and ventral body flexions that overrides all other
behaviors. We have explored the structure of the central pattern
generator (CPG) in the cerebropleural ganglion as part of a study of
neural network interactions underlying decision making in normal
behavior. The CPG comprises at least eight bilaterally paired
interneurons, each of which contributes and is phase-locked to the swim
rhythm. Dorsal flexion is mediated by hemiganglion ensembles of four
serotonin-immunoreactive neurons, the As1, As2, As3, and As4, and an
electrically coupled pair, the A1 and A10 cells. When stimulated, A10
commands fictive swimming in the isolated CNS and actual swimming
behavior in whole animals. As1-4 provide prolonged, neuromodulatory
excitation enhancing dorsal flexion bursts and swim cycle number.
Ventral flexion is mediated by the A3 cell and a ventral swim
interneuron, IVS, the soma of which is yet unlocated.
Initiation of a swim episode begins with persistent firing in A10,
followed by recruitment of As1-4 and A1 into dorsal flexion. Recurrent
excitation within the As1-4 ensemble and with A1/A10 may reinforce
coactivity. Synchrony among swim interneuron partners and bilateral
coordination is promoted by electrical coupling among the A1/A10 and
As4 pairs, and among unilateral As2-4, and reciprocal chemical
excitation between contralateral As1-4 groups. The switch from dorsal
to ventral flexion coincides with delayed recruitment of A3, which is
coupled electrically to A1, and with recurrent inhibition from
A3/IVS to A1/A10. The alternating phase relation may be
reinforced by reciprocal inhibition between As1-4 and IVS.
Pleurobranchaea's swim resembles that of the nudibranch
Tritonia; we find that the CPGs are similar in many details,
suggesting that the behavior and network are primitive characters
derived from a common pleurobranchid ancestor.
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