Abstract
ABSTRACTGrasping soft, irregular material is challenging both for animals and robots. The feeding systems of many animals have adapted to this challenge. In particular, the feeding system of the marine mollusk Aplysia californica, a generalist herbivore, allows it to grasp and ingest seaweeds of varying shape, texture and toughness. On the surface of the grasper of A. californica is a structure known as the radula, a thin flexible cartilaginous sheet with fine teeth. Previous in vitro studies suggested that intrinsic muscles, I7, are responsible for opening the radula. Lesioning I7 in vivo does not prevent animals from grasping and ingesting food. New in vitro studies demonstrate that a set of fine muscle fibers on the ventral surface of the radula – the sub-radular fibers (SRFs) – mediate opening movements even if the I7 muscles are absent. Both in vitro and in vivo lesions demonstrate that removing the SRFs leads to profound deficits in radular opening, and significantly reduces feeding efficiency. A theoretical biomechanical analysis of the actions of the SRFs suggests that they induce the radular surface to open around a central crease in the radular surface and to arch the radular surface, allowing it to softly conform to irregular material. A three-dimensional model of the radular surface, based on in vivo observations and magnetic resonance imaging of intact animals, provides support for the biomechanical analysis. These results suggest how a soft grasper can work during feeding, and suggest novel designs for artificial soft graspers.
Highlights
Aplysia californica is a generalist herbivore that feeds on red, green and brown seaweeds (Winkler and Dawson, 1963; Kupfermann and Carew, 1974; Carefoot, 1987), which differ biomechanically
In vitro experiments on I7 identified a new source of opening movements: sub-radular fibers (SRFs) In vitro experiments suggested that the I7 muscles were responsible for radular opening (Evans et al, 1996)
The reduced preparation in which the I7 muscles were studied consisted of the radular surface dissected free from the underlying musculature
Summary
Aplysia californica is a generalist herbivore that feeds on red, green and brown seaweeds (Winkler and Dawson, 1963; Kupfermann and Carew, 1974; Carefoot, 1987), which differ biomechanically. The red seaweeds Laurencia pacifica and Gigartina canaliculata have fernlike branching blades. The brown seaweed Macrocystis pyrifera (giant kelp) has blades that form large, leaf-like structures (laminae) supported by air-filled bladders. The green seaweed Ulva lactuca (sea lettuce) forms large sheets. Unlike snails that rasp algae, A. californica’s feeding apparatus must grasp and tear off pieces of seaweed whose shape, toughness and texture vary significantly, even within a single seaweed (Howells, 1942; Carefoot, 1987).
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