Neuronal Mechanisms for the Control of Body Orientation inClioneII. Modifications in the Activity of Postural Control System

The marine mollusk Clione limacina, when swimming, can stabilize different body orientations in the gravitational field. The stabilization is based on the reflexes initiated by activation of the statocyst receptor cells and mediated by the cerebro-pedal interneurons that produce excitation of the motoneurons of the effector organs; tail and wings. Here we describe changes in the reflex pathways underlying different modes of postural activity; the maintenance of the head-up orientation at low temperature, the maintenance of the head-down orientation at higher temperature, and a complete inactivation of the postural mechanisms during defense reaction. Experiments were performed on the CNS-statocyst preparation. Spike discharges in the axons of different types of neurons were recorded extracellularly while the preparation was rotated in space through 360° in different planes. We characterized the spatial zones of activity of the tail and wing motoneurons and the CPB3 interneurons mediating the effects of statocyst receptor cells on the tail motoneurons. This was done at different temperatures (10 and 20°C). The “fictive” defense reaction was evoked by electrical stimulation of the head nerve. At 10°C, a tilt of the preparation evoked activation in the tail motoneurons and wing retractor motoneurons contralateral to the tilt and in the wing locomotor motoneurons ipsilateral to the tilt. At 20°C, the responses in the tail motoneurons and in the wing retractor motoneurons occurred reversed; these neurons were now activated with the ipsilateral tilt. In the wing locomotor motoneurons the responses at 20°C were suppressed. During the defense reaction, gravitational responses in all neuron types were suppressed. Changes in the chains of tail reflexes most likely occurred at the level of connections from the statocyst receptor cells to the CPB3 interneurons. The changes in gravitational reflexes revealed in the present study are sufficient to explain the corresponding modifications of the postural behavior in Clione.