Biological spacetime and the temporal integration of functional modules:: A case study of dento-gnathic developmental timing

For the individual, coordination between tooth and jaw development is important to proper food acquisition and ingestion later in life. Among and within species, variation in dental and gnathic size, shape, and, in the case of teeth, number, must be mutually accommodating and functionally compatible. For these reasons, the development and evolution of these two systems should be closely integrated. Furthermore, the timing of dental development correlates tightly with life history events such as weaning. This correlation hints at a central regulation of the developmental timing of multiple systems that have tandem effects on physiology and behaviour. Important work on embryonic oral development continues to tease apart the molecular mechanisms that pattern jaw identity and establish tooth morphology and position in the alveolar bone. Still very poorly understood is what underlies rates and periods of gene activity such that pre- and postnatal tooth and jaw development are coordinated. Recent literature suggests at least some level of autonomy between permanent tooth and mandibular ontogenetic timing. However, whether the timing of these various signaling pathways is directly regulated or is an outcome of the pathways themselves is untested. Here, we review what is currently known about the embryonic signaling pathways that regulate tooth and jaw development in the context of time rather than space, as has been traditional. We hypothesize that the timing of mandibular and dental development is not directly mediated by a common factor but is an indirect outcome of strong selection for coordinated molecular pathways and growth trajectories. The mandible and lower jaw dentition is a powerful model with which to investigate the mechanisms that facilitate morphological change-in this case, the development and evolution-of organs that are closely integrated in terms of function, space and time.