Understanding the unique flowering sequence in Dipsacus fullonum: Evidence from geometrical changes during head development.

The genus Dipsacus is characterized by a remarkable bidirectional flowering sequence and a rare phyllotactic pattern. Considering that flower initiation and flowering sequence may be interconnected, we document the development of the head meristem in Dipsacus fullonum. Our results indicate a gradual change in the geometry of the head meristem beginning with a dome shaped stage, continuing with a remarkable widening in the middle part of the head meristem and ending in a spindle-like form. Quantitative data confirm that meristem expansion is higher in the middle part than at the base of the meristem. Likewise, the size of the flower primordia in the middle part of the young head is significantly larger than at the base soon after initiation. We conclude that the change in the geometry of the meristem and the availability of newly generated space result in the promotion of the middle flowers and the bidirectional flowering sequence at anthesis. Our investigation on phyllotactic patterns reveals a high tendency (30%) of the head meristem to insert or lose parastichies. This finding can also be attributed to changes in the expansion rate of the meristem. Dependent on the spatio-temporal relation between meristem expansion and primordia initiation, either flower primordia are promoted or additional parastichies appear. Our results emphasize the important role of geometry in flower development and phyllotactic pattern formation.