The radular apparatus of cephalopods

Abstract

This paper describes the ontogeny, breakdown and absorption of the radular teeth of cephalopods and, for the first time, considers the function of the ‘bolsters’ or radular support muscles.The radular ribbon, which bears many regularly arranged transverse rows of teeth one behind the other, lies in a radular canal that emerges from the radular sac. Here the radular teeth are formed by a set of elongate cells with microvilli, the odontoblasts. These are organized into two layers, the outer producing the radular membrane and the bases of the teeth, the inner producing the cusps. The odontoblasts also secrete the hyaline shield and the teeth on the lateral buccal palps, when these are present. At the front end of the radular ribbon the teeth become worn in feeding and are replaced from behind by new ones formed continuously in the radular sac, so that the whole ribbon moves forward during ontogeny. Removal of the old teeth is achieved by cells in the radular organs; these cells, which are formed from modified odontoblasts (‘odontoclasts’), dissolve the teeth and membranes and absorb them. There is a subradular organ in all cephalopods. InOctopus vulgaris, which bores into mollusc shells and crustacean carapaces, it is especially well–developed and there is also a supraradular organ.A characteristic feature of the cephalopod radular apparatus is the pair of large radular support muscles or ‘bolsters’. Their function seems never to have been investigated, but experiments reported here show that when they elongate, the radular teeth become erect at the bending plane and splayed, presumably enhancing their ability to rake food particles into the pharynx. The bolsters ofOctopusfunction as muscular hydrostats: because their volume is fixed, contraction of their powerful transverse muscles causes them to elongate. In decapods and in nautiloids each bolster contains a ‘support rod’ of semi–fluid material, as well as massive transverse musculature. This rod may elongate to erect the radular teeth. At the extreme front end of the bolsters inOctopusthere are many nerve fibres that may constitute a receptor organ signalling the movements of the radula against hard material. Such nerves are absent from decapods and from octopods that do not bore holes.The buccal mass ofNautilusis massive, with heavily calcified tips to the beaks and a wide radular ribbon, with 13 rather than nine elements in each row. Nevertheless all the usual coleoid features are present in the radular apparatus and the teeth are formed and broken down in the same way. However,Nautilushas a unique structure, the radular appendage. This comprises a papillate mass extending over the palate in the mid–line and forming paired lateral masses that are in part secretory. The organ is attached to the front of the radula by muscles and connective tissue. Its function is unknown.