Tinkering with transcription factor networks for developmental robustness of Ranunculales flowers

The flowers of core eudicots and monocots are generally determined by the number of floral organs they produce, and their developmental set-up tolerates little change from the bauplan once the floral primordium is initiated. Many species outside the core eudicots and monocots are more plastic in the number of floral organs they produce. For example, the Nymphaeales (water lilies), within the basal angiosperms, arrange their floral organs spirally and show smooth transitions between floral organs, and many Ranunculales (buttercups) produce variable numbers of stamens by adjusting the number of stamen whorls generated from a specialized ring meristem. However, the interactions of regulatory genes governing those processes are unknown.

This review provides an overview of the functional analyses of floral homeotic genes carried out in Ranunculales, summarizing knockdown and mutant phenotypes, and protein interactions to identify similarities and differences within the Ranunculales and in comparison with core eudicots. Floral gene regulatory networks in Ranunculales are identified showing intensive re-wiring amongst the floral homeotic genes to allow some degree of plasticity. The 'fading-border' model of floral organ identity evolution is extended by a hypothesis on how developmental plasticity can be achieved by interdependent regulation of floral homeotic genes. One aspect of floral plasticity may be achieved by regulation of the activity of a stamen-generating ring meristem and first ideas on its control are presented. While the amazing conservation of the major floral organ identity programme is being unravelled by analysing floral homeotic gene function and expression, we are only just beginning to understand the evolution of the gene network governing the organ identity genes, e.g. how plasticity can be achieved, and which aspects foster the robustness of the core eudicot floral bauplan.