The ultrastructure and ontogeny of pollen inHelleborus FoetidusL.: III. the formation of the pollen grain wall

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Abstract
The first recognizable elements of the pollen grain wall of Helleborus foetidus are initiated in the cellulosic primexine which is formed immediately outside the microspore cytoplasm while the pollen grains are still in the tetrad configuration and enveloped in a thick layer of callose. Elements of the primexine give rise to the precursors of the rod-like bacula of the mature exine. The bacula increase in electron density due to the rapid deposition of sporopollenin, and begin to expand laterally at the outer side to form the tectum. There follows a lateral expansion on the inner side to form the foot-layer. Further deposition of sporopollenin is continued and all elements of the pollen grain wall expand outwards and laterally as the pollen grain enlarges. The enveloping callose disappears and the pollen grains are free in the thecal cavity. A secondary exine is deposited below the primary exine, particularly around the furrows. Initially this process involves a number of thin electron-transparent lines or lamellae about 4 nm thick that appear to arise from the cytoplasm and provide a locus around which sporopollenin is deposited. As the deposition proceeds, the lamellae thicken and finally merge with each other to form the secondary exine. No sign of the lamellae can be seen in the mature pollen grain wall. Towards the end of secondary exine formation the deposition of sporopollenin does not appear to be centred on thin lamellae, but appears as small granules which gradually coalesce. The secondary exine remains discontinuous in the region of the furrow, but becomes consolidated in the inter-furrow regions. As the pollen grain matures the sporopollenin, which is electron-dense when initially deposited, becomes progressively less so. The final stage in development is the deposition of the cellulosic intine, which forms inside the secondary exine and is associated with increased dictyosome activity and randomly oriented microtubules.