Rapid shell closure in the brachiopodsTerebratulina retusaandTerebrataua transversa

Abstract

Rapid shell closure in articulate brachiopods, occurring by a twitch contraction of the the ‘quick’ adductor muscles, is a response to disturbance or to physiological requirements of the organism. The relative simplicity of the closing system permits a detailed analysis of the functional architecture of the mechanism and the underlying principles of skeleto-muscular organization, in terms of (1) basic kinematic properties of the system (speeds and times of closure), (2) hydrodynamic reactions resisting closure, and (3) considerations of muscle physiology and mechanics.Analyses of shell closure in the brachiopodsTerebratulina retusafrom the Firth of Lorn, Scotland, andTerebratalia transversafrom Puget Sound, USA, reveal (1) shell-closing times of the order of 50 to 70 ms, (2) closing velocities of the order of 3·5 radians s-1, from initial gapes of about 0·05 to 0·2 rad, and (3) muscle moment forces and hydrodynamic reactions with magnitudes of the order of 5 × 10-4N m (5 g cm). Muscle tensions developed in the ‘quick’ adductor muscle are of the order of 105N m2, and contraction velocities are of the order of one muscle length per second. Hydrodynamic reactions are a fundamental constraint on the closing mechanism, as determined by the concordance of actual closing events with predictions of a hydrodynamic model.