A Pair of Identified Interneurons inAplysiathat Are Involved in Multiple Behaviors Are Necessary and Sufficient for the Arterial-Shortening Component of a Local Withdrawal Reflex

Abstract

A bilateral pair of cerebral interneurons, called CC5, contribute to the generation of a number of different behaviors involving head movements. Each cell sends its axon to the ipsilateral and contralateral pedal and pleural ganglia. A weak tactile stimulus to the head excites the ipsilateral CC5; a strong stimulus excites both the ipsilateral and contralateral cells. Firing of CC5 produces powerful shortening of the ipsilateral pedal artery (PA) by means of monosynaptic excitation of the pedal artery shortener (PAS) neuron, the single motor neuron for the artery. A weak touch to a tentacle excites the ipsilateral PAS and evokes a local withdrawal response accompanied by shortening of the ipsilateral PA. In vivo recording of the pedal artery nerve (PAn) showed that PAS was activated bilaterally during defensive head withdrawal elicited by a strong stimulus and was activated unilaterally by a weak stimulus. The responses were eliminated by cutting the ipsilateral cerebral-pleural connective (C-PLC). Electrical stimulation of the cerebral-pleural connective provided evidence that all of the excitatory input to PAS via this connective is provided by CC5. A variety of experimental results indicates that during a local withdrawal reflex of the tentacle, CC5 is necessary and sufficient for the unilateral PA-shortening component of the response and therefore functions as a command neuron for a component of the behavior. The data suggest that during defensive head withdrawal, the two CC5 neurons may act conjointly as a two-neuron command system that is necessary and sufficient for the bilateral arterial-shortening component of the behavior.