12.3 Seed plants: gymnosperms

The first plants to colonize land were most likely closely related to modern-day mosses (bryophytes) and are thought to have appeared about 500 million years ago. They were followed by liverworts (also bryophytes) and primitive vascular plants, the pterophytes, from which modern ferns are derived. The life cycle of bryophytes and pterophytes is characterized by the alternation of generations. The completion of the life cycle requires water, as the male gametes must swim to the female gametes. The male gametophyte releases sperm, which must swim—propelled by their flagella—to reach and fertilize the female gamete or egg. After fertilization, the zygote matures and grows into a sporophyte, which in turn will form sporangia, or "spore vessels,” in which mother cells undergo meiosis and produce haploid spores. The release of spores in a suitable environment will lead to germination and a new generation of gametophytes.

The evolution of seed plants

In seed plants, the evolutionary trend led to a dominant sporophyte generation, in which the larger and more ecologically significant generation for a species is the diploid plant. At the same time, the trend led to a reduction in the size of the gametophyte, from a conspicuous structure to a microscopic cluster of cells enclosed in the tissues of the sporophyte. Lower vascular plants, such as club mosses and ferns, are mostly homosporous (produce only one type of spore). In contrast, all seed plants, or spermatophytes, are heterosporous, forming two types of spores: megaspores (female) and microspores (male). Megaspores develop into female gametophytes that produce eggs, and microspores mature into male gametophytes that generate sperm. Because the gametophytes mature within the spores, they are not free-living, as are the gametophytes of other seedless vascular plants. Heterosporous seedless plants are seen as the evolutionary forerunners of seed plants.

Seeds and pollen—two adaptations to drought—distinguish seed plants from other (seedless) vascular plants. Both adaptations were critical to the colonization of land. Fossils place the earliest distinct seed plants at about 350 million years ago. The earliest reliable record of gymnosperms dates their appearance to the Carboniferous period (359–299 million years ago). Gymnosperms were preceded by the progymnosperms (“first naked seed plants”). This was a transitional group of plants that superficially resembled conifers (“cone bearers”) because they produced wood from the secondary growth of the vascular tissues; however, they still reproduced like ferns, releasing spores to the environment. In the Mesozoic era (251–65.5 million years ago), gymnosperms dominated the landscape. Angiosperms took over by the middle of the Cretaceous period (145.5–65.5 million years ago) in the late Mesozoic era, and have since become the most abundant plant group in most terrestrial biomes.