Robustness and flexibility in nematode vulva development

The Caenorhabditis elegans vulva has served as a paradigm for how conserved developmental pathways, such as EGF-Ras-MAPK, Notch and Wnt signaling, participate in networks driving animal organogenesis. Here, we discuss an emerging direction in the field, which places vulva research in a quantitative and microevolutionary framework. The final vulval cell fate pattern is known to be robust to change, but only recently has the variation of vulval traits been measured under stochastic, environmental or genetic variation. Whereas the resulting cell fate pattern is invariant among rhabditid nematodes, recent studies indicate that the developmental system has accumulated cryptic variation, even among wild C. elegans isolates. Quantitative differences in the signaling network have emerged through experiments and modeling as the driving force behind cryptic variation in Caenorhabditis species. On a wider evolutionary scale, the establishment of new model species has informed about the presence of qualitative variation in vulval signaling pathways.