Axial Patterning Beyond the Individual: Colony-level Organization in a Siphonophore Colony

Colonial animals composed of clonally produced units can achieve a high degree of functional integration, challenging the distinction between an individual and a higher-order organism. Siphonophores (Cnidaria: Hydrozoa) exemplify this condition, forming highly organized colonies in which genetically identical zooids are specialized for functions such as locomotion, feeding, and reproduction, and are precisely arranged along a shared stem. All zooids arise from two spatially separated budding zones, the nectosomal and siphosomal growth zones, suggesting that positional information along the stem patterns colony organization at the level of the colony rather than individual zooids. However, the molecular basis of this colony-level axial patterning remains poorly understood. Here, we analyze gene expression along the stem of the siphonophore Agalma okenii using RNA sequencing and in situ hybridization chain reaction (HCR). We show that conserved developmental regulators, including Hox and Wnt pathway genes, exhibit region-specific expression corresponding to distinct budding zones and zooid distributions. These results indicate that canonical axial patterning systems are deployed at the level of the colony axis. Our findings demonstrate that developmental gene networks classically associated with anterior-posterior patterning can operate at a higher level of biological organization, providing a mechanistic framework for the evolution of integrated, superorganism-like body plans in colonial animals.