Metamorphosis of the marine bryozoan Membranipora membranacea: An ultrastructural study of rapid morphogenetic movements

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Abstract
The colonial marine bryozoan Membranipora membranacea produces a planktotrophic cyphonautes larva that is encased by a triangular bivalved shell. Following a relatively long free-swimming phase, the cyphonautes larva settles and undergoes a dramatic metamorphosis to become the sessile progenitor of the colony, referred to as the ancestrula. This paper examines the initial morphogenetic movements of metamorphosis that transform the cyphonautes larva into an incipient ancestrula. At the onset of metamorphosis, contractions of the striated median muscles situated along the anterior and posterior margins of the larva cause a retraction of the larval apical organ and a centripetal movement of the anterior and posterior ends of the larva. Concurrently, the ciliated corona at the base of the larva is pulled within the shell by contractions of the striated lateral muscles. As the larva assumes a more spherical shape, the posterior margins of the shell are spread apart, and the internal sac is everted. Eversion of the sac is apparently achieved by contractions of the lateral muscles that cause a buckling of the shell in the apical-basal direction. The neck region of the everted sac secretes adhesive granules that attach the larva to the substratum. Subsequently, contractions of the nonstriated sac muscles fold the valves of the shell over each other and draw the larva toward the substratum. The initial events of metamorphosis that culminate in the attachment and flattening of the larva are completed in 10–15 seconds. In the subsequent few minutes, the lateral edges of the everted sac fuse with the neighboring margins of the aboral epithelium underlying the shell and thus form the fully sealed body wall of the incipient ancestrula.