Mesoderm in spiralians: the organizer and the 4d cell

The Spiralia is a clade of protostome invertebrate phyla that share a highly conserved mode of early development. Spiralian development is characterized by regularities in the arrangement of early cleavages, the fates of the cells that are produced by these divisions, and the development of the distinctive trochophore larva. Because of the strong conservation in early development, homologies can be identified between cells in divergent taxa. Some of the most striking examples of conservation in the spiralian embryo are in the cells that generate the mesoderm. The specification of the mesodermal precursors has been well characterized by embryological approaches, and recently the molecular mechanisms of mesoderm specification are starting to be elucidated. This review examines the development of mesoderm in spiralians in a comparative context, with particular focus on the relationship between the mesendodermal cell 4d and the embryonic organizer.