Printed in Great Britain THE ULTRASTRUCTURE AND ONTOGENY OF POLLEN IN HELLEBORUS FOETIDUS L. IV. POLLEN GRAIN MATURATION

The final stages of Helleborus pollen-grain ontogeny, which culminate in maturation and germination of the grain, have been investigated at the ultrastructural level. Following the deposition of primary and secondary exine, and during the early stages of intine formation, the microspore passes through a vacuolate phase, in which the cytoplasm appears devoid of most organelles other than the prominent nucleus. The formation of the vacuole results in the displacement of the nucleus to one side of the pollen grain. The vacuole quickly disappears and a number of organelles reappear in the cytoplasm, in particular the dictyosomes and strands of endoplasmic reticulum, with associated grey bodies. Following mitotic division of the pollen grain, the first signs of the generative cell wall appear as a pair of tightly appressed unit membranes in the narrow strand of cytoplasm separating the two newly formed generative and vegetative nuclei. As development proceeds, the space between the two membranes gradually fills with an electron-transparent material similar to the substance found in the numerous dictyosome-derived vesicles which, together with the endoplasmic reticulum, are both closely associated with the developing cell wall. The generative cell wall fuses with the cellulosic intine, which has gradually increased in amount during these stages, and the cell division is complete