Phylogenetics of swimming behaviour in Medusozoa: the role of giant axons and their possible evolutionary origin

Although neural tissues in cnidarian hydroids have a nerve net structure, some cnidarian medusae contain well-defined nerve tracts. As an example, the hydrozoan medusa Aglantha digitale has neural feeding circuits that show an alignment and condensation, which is absent in its relatives Aequorea victoria and Clytia hemisphaerica. In some cases, neural condensations take the form of fast propagating giant axons concerned with escape or evasion. Such giant axons appear to have developed from the fusion of many, much finer units. Ribosomal DNA analysis has identified the lineage leading to giant axon-based escape swimming in Aglantha and other members of the Aglaura clade of trachymedusan jellyfish. The Aglaura, along with sister subclades that include species such as Colobonema sericeum, have the distinctive ability to perform dual swimming, i.e. to swim at either high or low speeds. However, the form of dual swimming exhibited by Colobonema differs both biomechanically and physiologically from that in Aglantha and is not giant axon based. Comparisons between the genomes of such closely related species might provide a means to determine the molecular basis of giant axon formation and other neural condensations. The molecular mechanism responsible may involve ‘fusogens’, small molecules possibly derived from viruses, which draw membranes together prior to fusion. Identifying these fusogen-based mechanisms using genome analysis may be hindered by the many changes in anatomy and physiology that followed giant axon evolution, but the genomic signal-to-noise ratio may be improved by examining the convergent evolution of giant axons in other hydrozoa, such as the subclass Siphonophora.