Branchial innervation and ciliary control in the ascidian Corella.

Abstract

The cilia lining the stigmata of the branchial sac of an ascidian circulate water through the animal. These stigmatal cilia are under nervous control; when either siphon is stimulated, both siphons close by muscular contractions and at the same time the stigmatal cilia stop beating simultaneously in all parts of the branchial sac. Spontaneous ciliary arrests may also occur, with or without associated closure of the siphons. Elements of the branchial nervous system that run in the gill bars are assumed to be concerned in coordination of the ciliary arrests. The majority of the branchial nerve fibres emerge dorsally from the visceral nerves that form the posterior brain roots, although nerves are also believed to enter the branchial sac along its anterior margin. No cell bodies could be found in the branchial nerves or in the visceral nerves, so that the cell bodies of the branchial nerve fibres are assumed to lie in the central nervous system. The branchial nerve fibres form a peripheral conducting net extending throughout the branchial sac. Branches of these nerve fibres terminate in contact with some of the ciliated cells; cell-to-cell conduction (through close junctions?) probably spreads excitation to the other ciliated cells. Nerve-nerve junctions appear to be more sensitive to curare than those between nerves and ciliated cells. Electrical recordings from the branchial sac, obtained with suction electrodes, show that arrest of the cilia is accompanied by electrical activity, and that prolonged arrest is maintained by trains of regular pulses. Intracellular microelectrodes in the ciliated cells indicate that these cells have a negative resting potential of 30-40 mV, and that a ciliary arrest is associated with a positive-going spike of 45-50 mV. The externally recorded ‘ciliary arrest potentials’ probably represent the coordinated depolarization of many ciliated cells. The rhythmical character of the trains of pulses presumably depends on pacemaker activity; this is not localized, since intact organisms or isolated small portions of the branchial sac are capable of generating similar trains of pulses. During the arrest response the stigmatal cilia first perform a reverse beat, then maintain the reverse position for several seconds before slowly relaxing and after several more seconds recommencing to beat with progressively increasing amplitude. The duration of the arrest response varies in media with different concentrations of the common cations, and also varies in response to repetitive stimulation, in a manner which suggests that the depolarization of the ciliated cells is associated with an influx of Ca2+, so that the ciliary control here may have some close parallels with that described forParamecium.