Structure and mechanics of starfish body wall.

The structure of the dorsal body wall of the starfish Echinaster spinulosus was studied using polarized light microscopy of frozen tissues, scanning electron microscopy and histology. The collagen fibres of the body wall form a three-dimensional orthogonal web. Voids in the web contain ossicles and papulae. The orthogonal web delivers dimensional stability but allows shear necessary for ray torsion. The ossicles and fibres interact to load the fibres in tension and the ossicles in compression. Strain rates of the dorsal body wall were measured on live animals during typical movements. Uniaxial tension tests of the body wall yielded Young's moduli of 267 MPa (longitudinal), 249 MPa (transverse) and 353 MPa (bias); curves were essentially linear. The body wall was approximately linearly viscoelastic and showed hysteresis at 0.01 Hz. Stress relaxation over five decades of time (in seconds) yielded relaxation spectra with peaks in relaxation time at 2.96-3.35, depending on test direction. Stress relaxation caused the connective tissue to soften. The surface of fractured stress-relaxed tissue revealed wispy, dissociated fibril tufts, whereas unrelaxed fractures produced blunt-ended fibre bundles. Neural control was necessary for body wall integrity.