The word “desolate” in the passage is closest in meaning to
[#paragraph1]The evolutionary history of plants has been marked by a series of adaptations. The ancestors of plants were photosynthetic single-celled organisms probably similar to today’s algae. Like modern algae, the organisms that gave rise to plants [#highlight1]presumably[/highlight1] lacked true roots, stems, leaves, and complex reproductive structures such as flowers. All of these features appeared later in the evolutionary history of plants. Of today’s different groups of algae, green algae are probably the most similar to ancestral plants. This supposition stems from the close phylogenetic (natural evolutionary) relationship between the two groups. DNA comparisons have shown that green algae are plants’ closest living relatives. In addition, other lines of evidence support the hypothesis that land plants evolved from ancestral green algae: green algae used the same type of chlorophyll and accessory pigments in photosynthesis as do land plants. This would not be true of red or brown algae. Green algae store food as starch, as do land plants and have cell walls made of cellulose, similar in composition to those of land plants. Again, the food storage and cell wall molecules of red and brown algae are different.
[#paragraph2]Today green algae live mainly in freshwater, suggesting that their early evolutionary history may have occurred in freshwater habitats. If so, the green algae would have been [#highlight3]subjected to[/highlight3] environmental pressures that resulted in adaptations that enhanced their potential to give rise to land-dwelling organisms.
[#paragraph3][#insert1]The environmental conditions of freshwater habitats, unlike those of ocean habitats, are highly variable. [#insert2] Water temperature can fluctuate seasonally or even daily, and changing levels of rainfall can lead to fluctuations in the concentration of chemicals in the water or even to periods in which the aquatic habitat dries up. [#insert3] Ancient freshwater green algae must have evolved features that enabled them to withstand extremes of temperature and periods of dryness. [#insert4] These adaptations served their descendants well as they invaded land.
[#paragraph4]The terrestrial world is green now, but it did not start out that way. When plants first made the transition ashore more than 400 million years ago, the land was barren and [#highlight5]desolate[/highlight5], inhospitable to life. [#highlight6]From a plant’s evolutionary viewpoint, however, it was also a land of opportunity, free of competitors and predators and full of carbon dioxide and sunlight (the raw materials for photosynthesis, which are present in far higher concentrations in air than in water).[/highlight6] So once natural selection had shaped the adaptations that helped plants overcome the obstacles to terrestrial living, plants prospered and diversified.
[#paragraph5]When plants pioneered the land, they faced a range of challenges [#highlight8]posed[/highlight8] by terrestrial environments. On land, the supportive buoyancy of water is missing, the plant is no longer bathed in a nutrient solution, and the air tends to dry things out. These conditions favored the evolution of structures that support the body, vessels that transport water and nutrients to all parts of the plant, and structures that conserve water. The resulting adaptations to dry land include some structural features that arose early in plant evolution; now these features are common to virtually all land plants. They include roots or rootlike structures, a waxy cuticle that covers the surfaces of leaves and stems and limits the evaporation of water, and pores called stomata in leaves and stems that allow gas exchange but close when water is scarce, thus reducing water loss. Other adaptations occurred later in the transition to terrestrial life and are now widespread but not universal among plants. These include conducting vessels that transport water and minerals upward from the roots and that move photosynthetic products from the leaves to the rest of the plant body and the stiffening substance lignin, which supports the plant body, helping it expose maximum surface area to sunlight.
[#paragraph6]These adaptations allowed an increasing diversity of plant forms to exploit dry land. Life on land, however, also required new methods of transporting sperm to eggs. Unlike aquatic and marine forms, land plants cannot always rely on water currents to carry their sex cells and disperse their fertilized eggs. So the most successful groups of land plants are those that evolved methods of fertilized sex cell dispersal that are independent of water and structures that protect developing embryos from drying out. Protected embryos and waterless dispersal of sex cells were achieved with the origin of seed plants and the key evolutionary innovations that they introduced: pollen, seeds, and, later, flowers and fruits.