Establishing Bilateral Symmetry in Hydrozoan Planula Larvae, a Review of Siphonophore Early Development

Siphonophores are colonial hydrozoans, characterized by complex colony organization and unparalleled zooid functional specialization. Recent genomic studies have offered an evolutionary perspective on how this morphological complexity arose, but a molecular characterization of symmetry breaking in siphonophore embryonic development is still largely missing. Here, bringing together historical data on early development with new immunohistochemical data, we review the diversity of developmental trajectories that lead to the formation of bilaterally symmetric planula larvae in siphonophores. Embryonic development, up to the planula stage, is remarkably similar across siphonophore phylogeny. Then, with the appearance of the lateral endodermal thickening (= ventral endoderm), larval development diverges between taxa, differing in the location and patterning of the primary buds, chronology of budding, establishment of growth zones, and retention of larval zooids. Our work also uncovers a number of open questions in siphonophore development, including homology of different zooids, mechanisms underlying formation and maintenance of spatially restricted growth zone(s), and molecular factors establishing a secondary dorsal-ventral axis in planulae. By discussing siphonophore development and body axes within the broader cnidarian context, we then set the framework for future work on siphonophores, which is finally achievable with the advent of culturing methods.