The kinematics of swallowing in the buccal mass of Aplysia californica.

Changes in the positions, shapes and movements of the feeding apparatus (buccal mass) of the marine mollusc Aplysia californica were studied in intact, transilluminated juveniles. The buccal mass assumes characteristic shapes as its internal structure, the radula/odontophore, moves anteriorly (protracts) or posteriorly (retracts). These shapes are especially distinctive when the radula/odontophore has protracted forwards fully, is close to its resting or neutral position, or has retracted backwards fully. We refer to the shapes that occur at full protraction, transition and full retraction as shape 1 (spherical), shape 2 (ovoid) and shape 3 (gamma-shaped), respectively. We introduce this shape nomenclature in order to avoid confusion with the existing terms protraction and retraction, which we reserve exclusively to describe the direction of movement of the radula/odontophore. The observed shape changes do not agree with those predicted on the basis of in vitro observations of a feeding head preparation, but are similar to shapes observed in vitro in the snail Lymnaea stagnalis. The buccal mass also rotates approximately 10 degrees dorsally during retraction, pivoting on the attachment to the mouth, before the subsequent protraction and return of the buccal mass to the transition shape. This rotation may be due to activation of the extrinsic muscles of the buccal mass. Plots of the buccal mass shape parameters eccentricity versus ellipticity create a two-dimensional shape space, which accurately quantifies the subtle transitions of shape between the different phases of the feeding cycle. Quantitative differences are observed between pure swallows and swallows with tearing behavior, but the qualitative shapes are similar. Hysteresis in the shape space plots of most swallows provides evidence for the hypothesis that protraction and retraction each have distinct 'active' and 'return' phases. The observed kinematic pattern imposes constraints on the internal structures of the buccal mass and may be used to infer the shape and positions of the radula and odontophore.