Microspore development in Annona (Annonaceae): Differences between monad and tetrad pollen

Pags.- Figs.- Tabls. The definitive version is available at: http://www.amjbot.org/• Premise of the study: Permanent tetrads are the most common form of pollen aggregation in flowering plants. The production of pollen in monads is plesiomorphic in angiosperms, but the aggregation into tetrads has arisen independently different times during the evolution of flowering plants. The causes behind the recurrent evolution of pollen aggregation from monads remain elusive. Permanent tetrad pollen is quite common in the Annonaceae, the largest family in the early-divergent order Magnoliales. In some genera, such as Annona, both tetrad- and monad-producing species can be found. • Methods: In this comparative study of pollen development, we use immunolocalization, cytological characterization, and enzymatic assays of four species in the genus Annona and one species in its closely related genus Asimina that release pollen in tetrads and two species in the genus Annona that release pollen in monads. • Key results: The main difference between species with tetrad and monad pollen is a delayed digestion of callose and cellulose at the pollen aperture sites that resulted in nonlayering of the exine in these areas, followed by a rotation and binding of the young microspores at the aperture sites. • Conclusions: Small changes in development resulted in clear morphological changes on pollen dispersal time and open a window on the possible selective advantage of the production of aggregated pollen.This work was supported by Ministerio de Economía y Competitividad–European Regional Development Fund, European Union (AGL2010-15140, AGL2012-40239, and AGL2013-43732-R), the BBVA Foundation (BIOCON 08-184/09), INIA (RF2012-00010 and RFP2012-00016), and Gob. Aragón A-43. J.L. was supported by a Juan de la Cierva postdoctoral grant of the Spanish Ministry of Science and Innovation.Peer reviewe