Development of the paratympanic pneumatic system of Japanese quail.

Avian heads are characterized as having two extensive air-filled systems lined with epithelia; the paranasal and paratympanic sinuses. Many diverticula derived from the paratympanic sinus system are known to reticulate with each other to form a single merged pneumatic space within the adult braincase. However, the development of these complex branching and reticulating epithelia has not been examined in detail. In this study, we describe the comprehensive developmental pattern of the paratympanic sinus and its associated soft tissues in a model bird, Japanese quail (Coturnix japonica). The data are derived from three-dimensional reconstructions based on histological sections and soft tissue enhanced micro-CT data. Those data provide the foundation of the complex hierarchical developmental pattern of the paratympanic sinus system. Moreover, associations with other tissues help establish key morphologies that identify each pneumatic entity. This study clarifies the developmental relationships of the ventral portions of the paratympanic sinus system, the siphoneal diverticulum and marginal sinus, based on the ligaments associated with the Eustachian tube. In addition, detailed histological pneumatic morphologies reveal hitherto unknown epithelial diversity, which may be indicative of equally complex developmental processes. We use the pneumatization of the quadrate as an example to support a close relationship with vascular growth and pneumatic epithelia invasion into ossified bone. We confirm pneumatic diverticula never enter into cartilages, possibly due to the absence of vasculature in these tissues. Lastly, we use the concept of a morphogenetic tree as a tool to help present the complex developmental pattern of the paratympanic sinus system and apply it toward inferring pneumatic morphologies in a nonavian theropod braincase.