SUMMARY In vertebrates, motile cilia line airways, oviducts and ventricles. Invertebrate cilia often control feeding, swimming and crawling, or gliding. Yet control and coordination of ciliary beating remains poorly understood. Evidence from the nudibranch mollusc, Tritonia diomedea , suggests that locomotory ciliated epithelial cells may be under direct electrical control. Here we report that depolarization of ciliated pedal epithelial (CPE)cells increases ciliary beating frequency (CBF), and elicits CBF increases similar to those caused by dopamine and the neuropeptide, TPep-NLS. Further,four CBF stimulants (zero external Cl - , depolarization, dopamine and TPep-NLS) depend on a common mode of action, viz . Ca 2+ influx, possibly through voltage-gated Ca 2+ channels, and can be blocked by nifedipine. Ca 2+ influx alone, however, does not provide all the internal Ca 2+ necessary for CBF change. Ryanodine receptor(RyR) channel-gated internal stores are also necessary for CBF excitation. Caffeine can stimulate CBF and is sensitive to the presence of the RyR blocker dantrolene. Dantrolene also reduces CBF excitation induced by dopamine and TPep-NLS. Finally, W-7 and calmidazolium both block CBF excitation by caffeine and dopamine, and W-7 is effective at blocking TPep-NLS excitation. The effects of calmidazolium and W-7 suggest a role for Ca 2+ -calmodulin in regulating CBF, either directly or via Ca 2+ -calmodulin dependent kinases or phosphodiesterases. From these results we hypothesize dopamine and TPep-NLS induce depolarization-driven Ca 2+ influx and Ca 2+ release from internal stores that activates Ca 2+ -calmodulin, thereby increasing CBF.

SUMMARY Behavioral experiments have demonstrated that the marine mollusc Tritonia diomedea can use the Earth's magnetic field as an orientation cue. Little is known, however, about the neural mechanisms that underlie magnetic orientation behavior in this or any other animal. In previous studies, two neurons in the brain of Tritonia , known as LPd5 and RPd5, were shown to respond with enhanced electrical activity to changes in earth-strength magnetic fields. We report evidence that two additional neurons, known as LPd6 and RPd6, also respond with increases in electrical activity when the magnetic field around the animal is altered. Anatomical analyses revealed that prominent neurites from the Pd6 cells are located within two ipsilateral nerves, pedal nerves 1 and 2. These nerves extend to the periphery of the animal and innervate tissues of the anterior ipsilateral foot and body wall. Electrophysiological recordings demonstrated that action potentials generated by the Pd6 cells propagate from the central ganglia toward the periphery. These results imply that the Pd6 cells play an efferent role in the magnetic orientation circuitry. Given that these cells contain cilio-excitatory peptides and that Tritonia crawls using ciliary locomotion, the Pd6 neurons may control or modulate cilia used in crawling, turning, or both.