Dual origins of mesoderm in a basal spiralian: Cell lineage analyses in the polyclad turbellarian Hoploplana inquilina

Evolutionary modifications in the origins and organization of the mesoderm represent significant events in the diversification of metazoan body plans. Within the Spiralia, mesoderm comprises ectomesoderm, which typically is generated by derivatives of the second and third quartets of micromeres, and endomesoderm, which is formed from the fourth quartet micromere of the D quadrant (4d). It has been held that endomesoderm generates the majority of adult mesodermal derivatives, while larval mesoderm is formed primarily from ectomesoderm. The evolutionary history of these mesodermal sources could be clarified by examining basal members of the Spiralia such as the polyclad turbellarians whose embryos exhibit canonical quartet spiral cleavage. Using the fluorescent lineage tracer DiI, we show that larval mesoderm is derived from only two cells, one from the ventral embryonic quadrant (2b, the 'mesectoblast' cell), and the other from the dorsal quadrant (4d, the mesentoblast cell). We compare these results with mesodermal origins in other spiralian phyla and conclude that a dual origin of mesoderm is a primitive feature of spiralian development. We also argue that ectomesoderm and endomesoderm should not be considered as the exclusive precursors of larval and adult mesoderm, respectively.