As bryophytes began to colonize the terrestrial surface, they produced organic acids during metabolism that aided in the breakdown of the rocky substrate. When they died, their organic matter mixed with the weathered rock, forming the Earth’s earliest soils. Formerly abundant to the first photosynthesizers to become terrestrial, access to sunlight became competitive as bryophytes expanded. This led to selection for individuals that could lift themselves higher and transport water throughout their tissues. Eventually, this selection resulted in the evolution of vascular tissue -- pipes that could bring water up from the ground so that parts of the plant could be raised upward, and those parts raised upward could transport their photosynthates down to the lower parts of the plant. The cells in the xylem (water-transporting vascular tissue) contained lignin, the tough, decay-resistant compound that wood is made out of. This rigid molecule in the vascular tissue allowed for structural support, allowing plants to grow taller -- some over 100 feet! The vascular system also allowed for the specialization of organs: roots for water absorption, leaves for photosynthesis, and stems for structural support.
Seedless vascular plants (SVPs) also began to rely more on the sporophyte stage. The sporophyte became the larger, nutritionally independent stage of the life cycle. Branching sporophytes offered more sites for meiosis to occur, resulting in increased opportunities for variation, which could be interpreted as more options in an increasingly competitive environment. There are approximately 20,000 known extant species, most of which are ferns.
SVPs are considered to be a paraphyletic group of organisms, forming two distinct lineages: Ferns and Lycophytes. These two lineages share the following characteristics:
- Morphology: Sporophytes develop complex tissues, including lignified vascular tissue, true roots, stems, and leaves. Sporophytes are branched, producing many sporangia. Gametophytes are reduced and thalloid. In some groups, the gametophyte is subterranean and parasitizes mycorrhizal fungi for sugars.
- Life cycle: Alternation of generations. Sporophyte dominant: sporophytes still grow from the gametophyte, but are now photosynthetic and the larger, longer-lived phase of the life cycle.
- Ecology: Gametes are still dispersed in water, so moisture is still required for fertilization.
Selection Pressures and Drivers
- Competition for sunlight. To get access to sunlight, SVPs needed to grow taller than bryophytes. However, this presents a problem of distributing water around the plant body to prevent drying out. Seedless vascular plants solved this problem with the adaptation of lignified vascular tissue. The lignin in the secondary walls of sclerenchyma cells allowed SVPs the structural support to grow taller.
- The initial forming of soils. Before the bryophytes, terrestrial surfaces were primarily rocky. The first land plants would have contributed to the chemical weathering of these rocks by producing acids during metabolism. After death, these plants add their organic matter to these weathering rocks, beginning to form Earth’s early soils. Fungi were likely involved in this process, as there are genes in the bryophytes for mycorrhizal relationships.