Cholinergic motor control of sea urchin tube feet: evidence for chemical transmission without synapses

Abstract

Isolated tube feet of Strongylocentrotus franciscanus contract briefly when the outer epithelium is touched. Similar twitch-like contractions can be induced by electrical stimulation of the outer surface of the tube foot. These responses appear to be chemically mediated. The following evidence indicates that the transmitter substance may be acetylcholine (ACh): ACh causes muscle contraction. This effect and that of electrical stimuli is potentiated by anticholinesterase agents and is antagonized by cholinergic blocking agents. Anaesthesia with chloralhydrate or chloretone abolishes responsiveness to mechanical or electrical stimulation but not to ACh. Desensitization with carbachol prevents responses to ACh and to mechanical or electrical stimulation. There are no neuromuscular synapses and no axons can be detected which cross the connective tissue layer which separates the muscle fibres from the subepithelial nerve plexus. The latter is known to contain conspicuous amounts of ACh; nerve terminals containing clear vesicles invest the outer surface of the connective tissue layer. All evidence indicates that chemical transmission involves diffusion of ACh (released from activated nerve terminals) across this connective tissue layer which is around 5 micron thick in fully extended tube feet but may have a thickness of 20 or even 25 micron in less extended ones. Calculations based on equations describing transmitter diffusion prove the feasibility of such a mechanism.