26.1 Evolution of seed plants

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 lifecycle of bryophytes and pterophytes is characterized by the alternation of generations, like gymnosperms and angiosperms; what sets bryophytes and pterophytes apart from gymnosperms and angiosperms is their reproductive requirement for water. The completion of the bryophyte and pterophyte life cycle requires water because 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 the sporangia, mother cells undergo meiosis and produce the haploid spores. Release of spores in a suitable environment will lead to germination and a new generation of gametophytes.

In seed plants, the evolutionary trend led to a dominant sporophyte generation, and at the same time, a systematic reduction in the size of the gametophyte: from a conspicuous structure to a microscopic cluster of cells enclosed in the tissues of the sporophyte. Whereas lower vascular plants, such as club mosses and ferns, are mostly homosporous (produce only one type of spore), all seed plants, or spermatophytes , are heterosporous. They form 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 critical adaptations to drought, and to reproduction that doesn’t require water—distinguish seed plants from other (seedless) vascular plants. Both adaptations were required for the colonization of land begun by the bryophytes and their ancestors. Fossils place the earliest distinct seed plants at about 350 million years ago. The first reliable record of gymnosperms dates their appearance to the Pennsylvanian period, about 319 million years ago ( [link] ). Gymnosperms were preceded by progymnosperms , the first naked seed plants, which arose about 380 million years ago. Progymnosperms were 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 into the environment. Gymnosperms dominated the landscape in the early (Triassic) and middle (Jurassic) Mesozoic era. Angiosperms surpassed gymnosperms by the middle of the Cretaceous (about 100 million years ago) in the late Mesozoic era, and today are the most abundant plant group in most terrestrial biomes.