The evolution of gnathostome development: Insight from chondrichthyan embryology.

Chondrichthyans (cartilaginous fishes) represent one of the two lineages of gnathostomes, the other being the osteicthyans (bony fishes). Classical studies on chondrichthyan embryology have strongly impacted our views of vertebrate body plan evolution, while recent studies highlight oviparous chondrichthyans as emerging vertebrate model systems that are amenable to experimental embryological manipulation. Here, we review three particular areas of interest in the field of chondrichthyan developmental biology-gastrulation, neural development, and appendage patterning-and we discuss recent findings within a broader chondrichthyan-osteichthyan comparative framework. In some cases, comparative studies of chondrichthyan and osteichthyan development reveal conserved patterns of gene expression in common developmental contexts. Studies of chondrichthyan gastrulation reveal conserved patterns of developmental gene expression, despite highly divergent modes of mesendoderm internalization, while molecular characterization of chondrichthyan neurogenic placodes indicates a conservation of placode transcription factor expression across gnathostome phylogeny. In other cases, comparative studies of chondrichthyan and osteichthyan development yield evidence of shared patterning mechanisms functioning in different developmental contexts. This is exemplified by studies on the development of chondrichthyan appendages-paired fins, median fins, and gill rays. These have demonstrated that a retinoic acid-responsive Shh-expressing signaling center functions to pattern the endoskeleton of gnathostome paired fins and chondrichthyan gill rays, while expression patterns of Tbx18 and HoxD family members are shared by gnathostome paired fins and chondrichthyan median fins. These findings fuel novel hypotheses of developmental genetic homology, and demonstrate how comparative studies of gnathostome development can provide insight into the evolutionary processes that underlie morphological diversity.