The ultrastructure and ontogeny of pollen in Helleborus Foetidus l. II. pollen grain development through the callose special wall stage

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Abstract
During the early stages of microsporocyte ontogeny in Helleborus foetidus L. there is protoplasmic continuity between the cells of the tapetum and between the individual sporogenous cells, but not between the two tissues. The plasma canals and plasmodesmata are progressively sealed off by the deposition of thick callose walls, so that by the first meiotic division, each pollen mother cell is isolated from its neighbours and from the surrounding tapetum. Callose is formed by dictyosomes in the individual pollen mother cells. The four meiocytes are separated by the deposition and coalescence of masses of callose forming in the cell plate area. The exine pattern is initiated at the surface of the young microspores while they are still invested with a thick wall of callose. Periclinally arranged endoplasmic reticulum lying just below the microspore cell membrane corresponds with the position of the furrows. The cell membrane in the interfurrow region thickens and becomes highly convoluted. A fibrous layer appears between the outer part of the convolutions and the callose, and locally it becomes less electron-dense at places that become filled with material of moderate electron density corresponding to the probacula; these in turn will become the bacula of the mature exine. In spite of an extensive examination of material prepared by a variety of techniques, no organelle or cytoplasmic component may be consistently associated with the positioning of the first signs of exine patterning.