Structural basis of harmomegathy: evidence from Boraginaceae pollen

Upon release from the anther, pollen grains can be exposed to dry environment and dehydrate. To survive in dry conditions, the pollen wall possesses the ability to fold itself due to water loss-harmomegathic mechanism. Apertures seem to function as the primary elements of harmomegathy as they are more elastic than the remainder of the pollen wall. Contribution of other sporoderm structures, surface features, and pseudocolpi in harmomegathy are usually not considered in palynological studies. The nature of pseudocolpi has not been properly understood until now, partly because of common use of acetolysis method as a standard procedure. Different structures involved in the harmomegathy mechanism were studied in Cryptantha celosioides, Cryptantha coryi, Heliotropium europaeum, Myosotis palustris, Rindera bungei, and Rindera tetraspis. Scanning electron microscopy was used to study harmomegathy in hydrated and dehydrated pollen grains. In addition, transmission electron microscopy was used to elucidate the ultrastructural basis of pseudocolpi and other harmomegathic structures with special attention to intine structure. Our data reveal that additional flexibility of the pollen wall in Boraginaceae is provided by pseudocolpi, rugulate surface, tectate-columellate ultrastructure, and a transverse groove. Curious triangular polar poroid areas are described in M. palustris.