Operant conditioning of head-waving in Aplysia. II. Contingent modification of electromyographic activity in identified muscles

Aplysia can readily exhibit operant conditioning of their head-waving response when bright light is used as aversive reinforcement (Cook and Carew, 1986). In the first paper of this series (Cook and Carew, 1989a), we showed that the electromyographic (EMG) activity of a discrete band of neck muscles, the lateral columellar muscles (LCMs) of Aplysia is significantly correlated with the component of head-waving (the horizontal component) that is modified during operant conditioning. In the present paper, we asked whether the EMG activity of the LCMs themselves could also be contingently modified, using the same procedures that produce operant conditioning of the behavioral response. Differential EMG from the LCMs was recorded in freely behaving animals with chronically implanted muscle cuff electrodes. Animals receiving aversive reinforcement (bright light) that was contingent upon specific patterns of LCM activity readily learned to alter their differential EMG output. Like operant conditioning of the head-waving response, this operant modification of LCM activity was rapidly acquired and was specific to the contingencies of reinforcement. These results show that a restricted group of muscles, the LCMs, exhibit the essential features of the head-waving system observed at the behavioral level: (1) their activity is significantly correlated with head-waving behavior, and (2) the LCMs are capable of operant modification of their output. Thus, this restricted response system provides a useful preparation for examining the neural mechanisms of operant conditioning of head-waving in Aplysia.