Screwed up: Spirality of segments and other iterated structures suggest an underlying principle of seriality in bilaterians.

This review deals with helicomery, that is, the specific malformation of a spiral arrangement of segments and other serial structures. Helicomery was first described in annelid and arthropod body segments. However, corresponding patterns occur in arthropod appendages and other bilaterians with serially arranged body parts, such as tapeworms, nematodes, vertebrates, and probably chitons. The specifics of the spirals such as length, orientation, and handedness are described. Most spirals are dorsal and comprise only a few loops. Helicomery is formed by a shift of cells during development or in adults caused by changes in cell adhesion or mechanical impacts such as lesions. A model for the formation of helicomery is proposed, which is based on medieval church labyrinths. These complex spiral structures are derived from concentric lines by the shift of relatively few tiles. This principle of "small causes, great effect" also applies to "spiral segments," because helicomery dissolves segmental patterns and questions the concept of segments as distinct structures. The widespread occurrence of helicomery in nonhomologous serial structures might indirectly indicate an underlying principle of seriality among Bilateria.