Factors Controlling the Change of Shape of Certain Nemertean and Turbellarian Worms

Abstract

1. Nemerteans and turbellarians have an inextensible fibre system around them in the form of a lattice of left- and right-handed spirals. The effect of this system on the change of shape on these worms has been analysed theoretically and compared with the observed behaviour of nine species of turbellarian and nemertean from widely differing habitats. 2. The following theoretical relationships have been studied: (a) Variation of the angle between the geodesics and the longitudinal axis of the worm during changes in length, and the role of the fibre system in limiting changes in length of the animal. (b) The change in cross-sectional shape during changes in length. (c) The extension of the fibres and the extensibility of the worms, assuming the fibres of the lattice to be elastic. 3. The species investigated conform with the theoretical predictions to varying degrees and have been grouped accordingly: (a) Geonemertes dendyi and Rhynchodemus bilineatus have low extensibilities and fit the prediction well. They are nearly circular in cross-section at all lengths as a result of their low extensibility and this is related to their terrestrial habit and need for water conservation. (b) Amphiporus lactifloreus, Lineus gesserensis and L. longissimus are moderately flattened in the relaxed position and have extensibilities between 6 and 10. They are marine crawling forms using cilia for locomotion and so must present a fairly large ciliated surface to the substratum. The fibre system does not limit contraction; the compression of the epithelial cells causes the observed extensibilities to fall a little short of the theoretical values. (c) Cerebratulus lacteus, Malacobdella grossa, Polycelis nigra and Dendrocoelum lacteum are very flattened forms and have very high theoretical extensibilities, but very low observed ones. The factors causing this are the thickness of the body-wall musculature (Cerebratulus), the limiting effect of longitudinal and circular reticulin fibres in the muscle layers, and the presence of dorso-ventral and diagonal muscles. Their flattened form is correlated with ecological factors (with swimming in Cerebratulus, with its parasitic life in the mantle of bivalves in Melacobdella) or with physical ones in turbellarians where a permanently flattened form is necessary for these worms to move by ciliary action.