Male Gametophyte Development and Evolution in Extant Gymnosperms

The male gametophytes of gymnosperms are characterized by the diversities in pollen morphology, cellular composition and pattern of cell division, pollen tube morphology, sperm delivery, growth pattern through the ovule and nucellus, and pollen tube wall composition both within and among the four living orders, i.e., Cycadales, Ginkgoales, Coniferales, and Gnetales. At dehiscence, gymnosperm pollen grains contain a variable number of cells yet none have sperm at this stage. Pollen germination in the ovule usually occurs within a few hours or days in gnetophytes, about a week or so in conifers and Ginkgo, or after several months in cycads. Complete development of the male gametophytes typically involves two to five mitotic divisions. Evolution of the male gametophyte appears to have involved a reduction of its component cells with prothallial cells being among those reduced or eliminated. There is a shift in the site of sperm discharge from a proximal position in pollen grains of cycads and Ginkgo to distal in conifers and gnetophytes. Two methods of sperm delivery occur in gymnosperms: zooidogamy, defined by pollen tubes with motile sperm as exhibited in cycads and Ginkgo, and siphonogamy, defined by pollen tubes with non-motile sperm which are directly delivered into the egg as exhibited in conifers and gnetophytes. Different pollen tube morphologies occur in the nucellus, i.e., branched and haustorial in cycads and Ginkgo, and unbranched and non-haustorial in conifers and gnetophytes. Pollen tubes form heterotrophic relationships with the nucellus, but it is only in cycads that intracellular penetration results in significant destruction of the nucellus. Pollen tube walls of gymnosperms contain cellulose and arabinogalactan proteins; however, pectins are prevalent in cycads and mixed -glucan in Ginkgo. A standard terminology to describe the cellular composition of the male gametophytes in gymnosperms is proposed. _____________________________________________________________________________________________________________