Stamen development in the Ericaceae. I. Anther wall, microsporogenesis, inversion, and appendages

Development of the introrse, tetrasporangiate, and normally dorsifixed and poricidal stamens has been studied at the gross morphological and cellular level in ten species of Ericaceae. Microsporogenesis, followed in four species, is normal, with cytokinesis simultaneous, forming tetrahedral tetrads. The tricolp(or)ate pollen is shed as permanent tetrads with each segment two-celled except in Enkianthus in which pollen grains are three-celled monads. Anther-wall development is similar in all four species initially, but no regular pattern of wall development could be recognized thereafter. The tapetum, of parietal origin, is binucleate, glandular, and mainly uniseriate. Viscin threads occur with the tetrads in the three rhododendroid species. A well-developed endothecium appears only in Enkianthus.Soon after stamen initiation, anthers of nine species invert at the eventual filament-anther junction to become introrse; in Enkianthus inversion occurs close to anthesis. Microsporogenesis starts during early inversion; greater cell elongation on the abaxial side of the young anther completes inversion by the late sporogenous-tissue stage. In Erica and, to a lesser extent Calluna, inversion results from greater abaxial than adaxial increase in cell number and length just above the filament-anther junction. The single vascular strand reflects the degree of inversion. Stamens of six species are appendaged; three have only awns, two only spurs, while one has both. Appendages arise from residual meristems after inversion is completed (or almost so) in all except Enkianthus. Awns develop at what will be the apex at maturity of each anther half. Their length and orientation vary among species. Only in Vaccinium do the awns become hollow (tubules). Spurs, varying in length, shape, and size, arise on the abaxial side from the filament, connective, or thecae.