Stereom morphogenesis and differentiation during regeneration of adambulacral spines of Asterias rubens (Echinodermata, Asteroida)

Summary

The very first mineral deposits appearing in regenerating fractured adambulacral spines of Asterias rubens are minute polyhedrons that cover the surface of fractured trabeculae. Polyhedrons fuse together forming a fold from which a microspine differentiates. Microspines develop into long linear trabeculae which send out lateral processes at regular length intervals. Lateral processes from adjacent trabeculae fuse together, bridging the trabeculae and giving the regenerate the typical meshwork structure of stereom. Most of the regenerate is built up according to this growth pattern which ensures its longitudinal growth. Simultaneously, the initial fascicular stereom of the stub sends out short radial processes which branch into upward and downward directed subprocesses. The latter fuse with their equivalents located above or below, building up longitudinal rows of stereom meshes. These rows then bridge together by additional branched or unbranched lateral processes, so forming a new stereom layer which progressively covers the whole stub. Up to three new layers of stereom are formed in this way at the stub periphery. These become continuous with the stereom layers of the regenerate by fusion of reciprocal subprocesses, so ensuring the continuity between the stub and the regenerate. In both structures the first stage of mineralization results in an open stereom. Stereom thickening occurs in a second stage of mineralization (that is chronologically separated from the formation of the open stereom) and results in the differentiation of the original stereom fabrics (i.e. fascicular stereom). Regeneration of removed spines starts with the formation of a new spine base made of labyrinthic stereom. The development of the latter mostly relies on short branched and unbranched processes which fuse with each other or with predifferentiated meshes. After completion of its base, the regenerating spine lengthens and thickens similarly to the regenerating fractured spines. The diversity of the stereom growth processes observed in the present work may be reduced to the combination of one to three elementary events, viz. the development of long linear processes, of short unbranched processes and of short branched processes. A survey of the literature allows the suggestion that the implementation of these elementary events is sufficient to describe most types of stereom morphogenesis.