Building an evolutionary innovation: differential growth in the modified vertebral elements of the zebrafish Weberian apparatus.

The Weberian apparatus, a complex assemblage of greatly modified vertebral elements, significantly enhances hearing within Otophysi. Ultimately we are interested in investigating the genetic mechanisms responsible for the origin, development and morphological diversification of these vertebral elements in the Weberian apparatus of otophysan fishes. However, a necessary first step involves identifying changes in growth of this region as compared with the vertebrae from which these modified elements purportedly derive. Using an ontogenetic series of the zebrafish, Danio rerio, we collected growth data for specific elements within the Weberian apparatus, including neural arches, ribs, and parapophyses. These data are compared to both serially homologous structures in posterior thoracic vertebrae (which act as internal controls) and vertebral elements from the same axial levels in three other non-otophysan teleosts. Significant differences in growth rate were found among serially homologous structures, as well as at equivalent axial levels in different species. Uniform changes in growth rates (in which all structures derived from a specific somite were equally affected) were not found, suggesting precise targeting of morphological change to specific structures. The variation in growth of anterior vertebrae in and among species was greater than expected. This variation in growth rates created developmental patterns unique to each species. Such patterns of growth may help illuminate the specific heterochronic mechanisms required for the origin and subsequent morphological diversification of the Weberian apparatus. This morphological diversity is exemplified by the multitude of forms seen in the cypriniform Weberian apparatus. Understanding patterns of growth in discrete elements of the Weberian apparatus allows us to hypothesize as to the specific developmental changes, likely constituting differences in gene expression in pathways involved in bone and cartilage differentiation, responsible for this morphological diversity.