Early Plant Life

Plant Adaptations to Life on Land

As organisms adapted to life on land, they had to contend with several challenges in the terrestrial environment. Water has been described as “the stuff of life.” The cell’s interior is a watery soup: in this medium, most small molecules dissolve and diffuse, and the majority of the chemical reactions of metabolism take place. Desiccation, or drying out, is a constant danger for an organism exposed to air. Even when parts of a plant are close to a source of water, the aerial structures are likely to dry out. Water also provides buoyancy to organisms. On land, plants need to develop structural support in a medium that does not give the same lift as water. The organism is also subject to bombardment by mutagenic radiation, because air does not filter out ultraviolet rays of sunlight. Additionally, the male gametes must reach the female gametes using new strategies, because swimming is no longer possible. Therefore, both gametes and zygotes must be protected from desiccation. The successful land plants developed strategies to deal with all of these challenges. Not all adaptations appeared at once. Some species never moved very far from the aquatic environment, whereas others went on to conquer the driest environments on Earth.

To balance these survival challenges, life on land offers several advantages. First, sunlight is abundant. Water acts as a filter, altering the spectral quality of light absorbed by the photosynthetic pigment chlorophyll. Second, carbon dioxide is more readily available in air than in water, since it diffuses faster in air. Third, land plants evolved before land animals; therefore, until dry land was colonized by animals, no predators threatened plant life. This situation changed as animals emerged from the water and fed on the abundant sources of nutrients in the established flora. In turn, plants developed strategies to deter predation: from spines and thorns to toxic chemicals.

Early land plants, like the early land animals, did not live very far from an abundant source of water and developed survival strategies to combat dryness. One of these strategies is called tolerance. Many mosses, for example, can dry out to a brown and brittle mat, but as soon as rain or a flood makes water available, mosses will absorb it and are restored to their healthy green appearance. Another strategy is to colonize environments with high humidity, where droughts are uncommon. Ferns, which are considered an early lineage of plants, thrive in damp and cool places such as the understory of temperate forests. Later, plants moved away from moist or aquatic environments using resistance to desiccation, rather than tolerance. These plants, like cacti, minimize the loss of water to such an extent they can survive in extremely dry environments.

The most successful adaptation solution was the development of new structures that gave plants the advantage when colonizing new and dry environments. Four major adaptations are found in all terrestrial plants: the alternation of generations, a sporangium in which the spores are formed, a gametangium that produces haploid cells, and apical meristem tissue in roots and shoots. The evolution of a waxy cuticle and a cell wall with lignin also contributed to the success of land plants. These adaptations are noticeably lacking in the closely related green algae—another reason for the debate over their placement in the plant kingdom.

Alternation of Generations

Alternation of generations describes a life cycle in which an organism has both haploid and diploid multicellular stages (Figure).

 The plant life cycle has haploid and diploid stages. The cycle begins when haploid (1n) spores undergo mitosis to form a multicellular gametophyte. The gametophyte produces gametes, two of which fuse to form a diploid zygote. The diploid (2n) zygote undergoes mitosis to form a multicellular sporophyte. Meiosis of cells in the sporophyte produces 1n spores, completing the cycle.
Alternation of generations between the 1n gametophyte and 2n sporophyte is shown. (credit: Peter Coxhead)

Haplontic refers to a lifecycle in which there is a dominant haploid stage, and diplontic refers to a lifecycle in which the diploid is the dominant life stage. Humans are diplontic. Most plants exhibit alternation of generations, which is described as haplodiplodontic: the haploid multicellular form, known as a gametophyte, is followed in the development sequence by a multicellular diploid organism: the sporophyte. The gametophyte gives rise to the gametes (reproductive cells) by mitosis. This can be the most obvious phase of the life cycle of the plant, as in the mosses, or it can occur in a microscopic structure, such as a pollen grain, in the higher plants (a common collective term for the vascular plants). The sporophyte stage is barely noticeable in lower plants (the collective term for the plant groups of mosses, liverworts, and lichens). Towering trees are the diplontic phase in the lifecycles of plants such as sequoias and pines.

Protection of the embryo is a major requirement for land plants. The vulnerable embryo must be sheltered from desiccation and other environmental hazards. In both seedless and seed plants, the female gametophyte provides protection and nutrients to the embryo as it develops into the new generation of sporophyte. This distinguishing feature of land plants gave the group its alternate name of embryophytes.

Sporangia in Seedless Plants

The sporophyte of seedless plants is diploid and results from syngamy (fusion) of two gametes. The sporophyte bears the sporangia (singular, sporangium): organs that first appeared in the land plants. The term “sporangia” literally means “spore in a vessel,” as it is a reproductive sac that contains spores Figure. Inside the multicellular sporangia, the diploid sporocytes, or mother cells, produce haploid spores by meiosis, where the 2n chromosome number is reduced to 1n (note that many plant sporophytes are polyploid: for example, durum wheat is tetraploid, bread wheat is hexaploid, and some ferns are 1000-ploid). The spores are later released by the sporangia and disperse in the environment. Two different types of spores are produced in land plants, resulting in the separation of sexes at different points in the lifecycle. Seedless non-vascular plants produce only one kind of spore and are called homosporous. The gametophyte phase is dominant in these plants. After germinating from a spore, the resulting gametophyte produces both male and female gametangia, usually on the same individual. In contrast, heterosporous plants produce two morphologically different types of spores. The male spores are called microspores, because of their smaller size, and develop into the male gametophyte; the comparatively larger megaspores develop into the female gametophyte. Heterospory is observed in a few seedless vascular plants and in all seed plants.

Photo shows sporangia in seedless plant Bryum capillare.
Spore-producing sacs called sporangia grow at the ends of long, thin stalks in this photo of the moss Esporangios bryum. (credit: Javier Martin)

When the haploid spore germinates in a hospitable environment, it generates a multicellular gametophyte by mitosis. The gametophyte supports the zygote formed from the fusion of gametes and the resulting young sporophyte (vegetative form). The cycle then begins anew.

The spores of seedless plants are surrounded by thick cell walls containing a tough polymer known as sporopollenin. This complex substance is characterized by long chains of organic molecules related to fatty acids and carotenoids: hence the yellow color of most pollen. Sporopollenin is unusually resistant to chemical and biological degradation. In seed plants, which use pollen to transfer the male sperm to the female egg, the toughness of sporopollenin explains the existence of well-preserved pollen fossils. Sporopollenin was once thought to be an innovation of land plants; however, the green algae Coleochaetes forms spores that contain sporopollenin.

Gametangia in Seedless Plants

Gametangia (singular, gametangium) are structures observed on multicellular haploid gametophytes. In the gametangia, precursor cells give rise to gametes by mitosis. The male gametangium (antheridium) releases sperm. Many seedless plants produce sperm equipped with flagella that enable them to swim in a moist environment to the archegonia: the female gametangium. The embryo develops inside the archegonium as the sporophyte. Gametangia are prominent in seedless plants, but are very rarely found in seed plants.

Apical Meristems

Shoots and roots of plants increase in length through rapid cell division in a tissue called the apical meristem, which is a small zone of cells found at the shoot tip or root tip (Figure). The apical meristem is made of undifferentiated cells that continue to proliferate throughout the life of the plant. Meristematic cells give rise to all the specialized tissues of the organism. Elongation of the shoots and roots allows a plant to access additional space and resources: light in the case of the shoot, and water and minerals in the case of roots. A separate meristem, called the lateral meristem, produces cells that increase the diameter of tree trunks.

Illustration shows the tip of a root. The cells in the tip are smaller than the more mature cells further up.
Addition of new cells in a root occurs at the apical meristem. Subsequent enlargement of these cells causes the organ to grow and elongate. The root cap protects the fragile apical meristem as the root tip is pushed through the soil by cell elongation.
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