Control of development and motility in the spermatozoids of lower plants

The spermatozoids of lower plants have long been recognized as remarkably complex motile gametes. Spermatozoids differ markedly from the other gametophyte cells that surround or give rise to them. Their differentiation process involves the synthesis and assembly of a complex cytoskeleton and a motile apparatus that can be simple or complex, having as few as two to as many as thousands of ciliary axonemes. An important aspect of spermiogenesis involves the de novo synthesis of basal bodies in a cytoplasmic particle known as the blepharoplast: that is, the cells that produce spermatocytes do not contain centrioles. Thus, these cells provide an ideal system in which to study the formation of basal bodies. The cytoskeletons of spermatozoids from different organisms display a common architecture, with a multilayered structure (MLS) at the anterior end of the cell and a dorsally situated planar ribbon of crosslinked microtubules extending the length of the elongated gamete. The function of the MLS is not known, but it could be involved in cell-body elongation during development and in the control of ciliary motility in the mature gamete, particularly during chemotaxis. The application of modern techniques on these cells can shed light on long-standing problems relating to spermiogenesis and motility.