Structure of staminate flowers, microsporogenesis, and microgametogenesis in Helosis cayennensis var. cayennensis (Balanophoraceae)

We analyzed the microgametogenesis and microsporogenesis of the male flowers of the holoparasitic Helosis cayennensis (Sw.) Spreng. var. cayennensis using optical and scanning electron microscopy. The unisexual flowers are embedded in a dense mass of uniseriate trichomes (filariae). Male flowers have a tubular 3-lobed perianth, with bilayered and non vascularized tepals. The androecium consists of three stamens with filaments and thecae connated into a synandrium. It has adnate a free central pistillode without megagametophyte. Staminal filaments, fused at their base to the perianth tube and distally free along a short section, have a single vascular bundle. The distal portion of the synandrium is formed by nine pollen sacs: six outer sacs are located laterally to each filament and three longer inner sacs. The anther wall consists of the epidermis, two parietal layers (that collapse at anther maturity), and an uninucleate secretory tapetum. There is no endothecium. During microsporogenesis, the stem cells produce tetrads of microspores by meiosis. The cytokinesis is simultaneous, forming tetrahedrally arranged tetrads. When pollen grains are in the tricellular state, the synandrium emerges from the mass of filariae, and anthers dehiscence occurs apically through longitudinal slits. In conclusion, despite the extreme reduction of flowers, the anatomic characteristics and gametophyte development of staminate flowers of H. cayennensis are perfectly normal and functional. They are thus highly similar to other genera of the holoparasitic subfamily Helosidoideae. Sterile parts of flowers and inflorescence maintain the same distinctive and aberrant features of the plant vegetative parts.