Dorsal-ventral patterning in amphioxus: current understanding, unresolved issues, and future directions.

How the embryonic body axis is generated is a fundamental question in developmental biology. The molecular mechanisms involved in this process have been the subject of intensive studies using traditional model organisms during the last few decades, and the results have provided crucial information for understanding the formation of animal body plans. In particular, studies exploring the molecular nature of Spemann's organizer have revealed the intricate interactions underlying several signaling pathways (namely the Wnt/β-catenin, Nodal and Bmp pathways) that pattern the dorsoventral (DV) axis in vertebrate embryos. Furthermore, recent comparative studies have shown that many of these signaling interactions are also employed in other non-vertebrate model organisms for their early embryonic axis patterning. These results suggest that there is deep homology in DV patterning mechanisms among bilaterian animals and that these mechanisms may be traced back to the common ancestor of cnidarians and bilaterians. However, the mechanism by which the DV axis became inverted in the chordate lineage relative to the DV axis in other bilaterian animals remains unclear. Cephalochordata (i.e., amphioxus) represent a basal chordate group which occupies a key phylogenetic position for explorations of the origin of the chordate body plan. In this review, we summarize what is currently known regarding the developmental mechanisms that establish the DV axis in amphioxus embryos. By comparing this to what is known in vertebrates, we can start to hypothesize about the ancestral DV patterning mechanisms in chordates and discuss their possible evolutionary origins.