6.1: Origins of Land Plants

Learning Objectives

By the end of this section, you will be able to do the following:

• Describe the traits shared by green algae and land plants
• Explain why charophytes are considered the closest algal relative to land plants
• Discuss the challenges to plant life on land
• Describe the adaptations that allowed plants to colonize the land
• Describe the timeline of plant evolution and the impact of land plants on other living things

 

The kingdom Plantae constitutes large and varied groups of organisms. There are more than 300,000 species of catalogued plants. Of these, more than 260,000 are seed plants. Mosses, ferns, conifers, and flowering plants are all members of the plant kingdom. Land plants arose within the Archaeplastida, which includes the red algae (Rhodophyta) and two groups of green algae, Chlorophyta and Charaphyta. Most biologists also consider at least some green algae to be plants, although others exclude all algae from the plant kingdom. The reason for this disagreement stems from the fact that only green algae, the Chlorophytes and Charophytes, share common characteristics with land plants (such as using chlorophyll a and b plus carotene in the same proportion as plants). These characteristics are absent from other types of algae.

 

Green Algae: Precursors to Land Plants

Until recently, all photosynthetic eukaryotes were classified as members of the kingdom Plantae. The brown and golden algae, however, are now reassigned to the protist supergroup Chromalveolata. This is because apart from their ability to capture light energy and fix CO2, they lack many structural and biochemical traits that are characteristic of plants. The plants are now classified, along with the red and green algae, in the protist supergroup Archaeplastida. Green algae contain the same carotenoids and chlorophyll a and b as land plants, whereas other algae have different accessory pigments and types of chlorophyll molecules in addition to chlorophyll a. Both green algae and land plants also store carbohydrates as starch. Their cells contain chloroplasts that display a dizzying variety of shapes, and their cell walls contain cellulose, as do land plants. Which of the green algae to include among the plants has not been phylogenetically resolved.

Green algae fall into two major groups, the chlorophytes and the charophytes. The chlorophytes include the genera Chlorella, Chlamydomonas, the “sea lettuce” Ulva, and the colonial alga Volvox. The charophytes include desmids, as well as the genera Spirogyra, Coleochaete, and Chara. There are familiar green algae in both groups. Some green algae are single cells, such as Chlamydomonas and desmids, which adds to the ambiguity of green algae classification, because plants are multicellular. Other green algae, like Volvox, form colonies, and some, like Ulva are multicellular (Figure 25.7). Spirogyra is a long filament of colonial cells. Most members of this genus live in fresh water, brackish water, seawater, or even in snow patches. A few green algae can survive on soil, provided it is covered by a thin film of moisture within which they can live. Periodic dry spells provide a selective advantage to algae that can survive water stress.

The chlorophytes and the charophytes differ in a few respects that, in addition to molecular analysis, place the land plants as a sister group of the charophytes. First, cells in charophytes and the land plants divide along cell plates called phragmoplasts, in which microtubules parallel to the spindle serve as guides for the vesicles of the forming cell plate. In the chlorophytes, the cell plate is organized by a phycoplast, in which the microtubules are perpendicular to the spindle. Second, only the charophytes and the land plants have plasmodesmata, or intercellular channels that allow the transfer of materials from cell to cell. In the chlorophytes, intercellular connections do not persist in mature multicellular forms. Finally, both charophytes and the land plants show apical growth—growth from the tips of the plant rather than throughout the plant body. Consequently, land plants and the charophytes are now part of a new monophyletic group called Streptophyta.

Figure 25.7 Green algae. Charophyta include (a) Spirogyra and (b) desmids. Chlorophyta include (c) Chlamydomonas, and (d) Ulva. Desmids and Chlamydomonas are single-celled organisms, Spirogyra forms chains of cells, and Ulva forms multicellular structures resembling leaves, although the cells are not differentiated as they are in higher plants (credit b: modification of work by Derek Keats; credit c: modification of work by Dartmouth Electron Microscope Facility, Dartmouth College; credit d: modification of work by Holger Krisp; scale-bar data from Matt Russell)

 

Evolution Connection

Algae and Evolutionary Paths to Photosynthesis: Some scientists consider all algae to be plants, while others assert that only the green algae belong in the kingdom Plantae. Still others include only the Charophytes among the plants. These divergent opinions are related to the different evolutionary paths to photosynthesis selected for in different types of algae. While all algae are photosynthetic—that is, they contain some form of a chloroplast—they didn’t all become photosynthetic via the same path.

The ancestors to the Archaeplastida became photosynthetic by forming an endosymbiotic relationship with a green, photosynthetic bacterium about 1.65 billion years ago. That algal line evolved into the red and green algae, and eventually into the modern mosses, ferns, gymnosperms, and angiosperms. Their evolutionary trajectory was relatively straight and monophyletic. In contrast, algae outside of the Archaeplastida, e.g., the brown and golden algae of the stramenopiles, and so on—all became photosynthetic by secondary, or even tertiary, endosymbiotic events; that is, they engulfed cells that already contained an endosymbiotic cyanobacterium. These latecomers to photosynthesis are parallels to the Archaeplastida in terms of autotrophy, but they did not expand to the same extent as the Archaeplastida, nor did they colonize the land.

Scientists who solely track evolutionary straight lines (that is, monophyly), consider only the Charophytes as plants. The common ancestor of Charophytes and land plants excludes the other members of the Archaeplastida. Charophytes also share other features with the land plants. These will be discussed in more detail in another section.

 

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Evolution of Land Plants

No discussion of the evolution of plants on land can be undertaken without a brief review of the timeline of the geological eras. The early era, known as the Paleozoic, is divided into six periods. It starts with the Cambrian period, followed by the Ordovician, Silurian, Devonian, Carboniferous, and Permian. The major event to mark the Ordovician, more than 500 million years ago, was the colonization of land by the ancestors of modern land plants. Fossilized cells, cuticles, and spores of early land plants have been dated as far back as the Ordovician period in the early Paleozoic era. The oldest-known vascular plants have been identified in deposits from the Devonian. One of the richest sources of information is the Rhynie chert, a sedimentary rock deposit found in Rhynie, Scotland (Figure 25.5), where embedded fossils of some of the earliest vascular plants have been identified.

Figure 25.5 Early vascular plant fossils. This Rhynie chert (a) contains fossilized material from vascular plants. Reconstruction of Cooksonia (b), the plant forms inside the circle. (credit b: modification of work by Peter Coxhead based on original image by “Smith609”/Wikimedia Commons; scale-bar data from Matt Russell)

 

Paleobotanists distinguish between extinct species, as fossils, and extant species, which are still living. The extinct vascular plants most probably lacked true leaves and roots and formed low vegetation mats similar in size to modern-day mosses, although some could reach one meter in height. The later genus Cooksonia, which flourished during the Silurian, has been extensively studied from well-preserved examples. Imprints of Cooksonia show slender branching stems ending in what appear to be sporangia. From the recovered specimens, it is not possible to establish for certain whether Cooksonia possessed vascular tissues. Fossils indicate that by the end of the Devonian period, ferns, horsetails, and seed plants populated the landscape, giving rising to trees and forests. This luxuriant vegetation helped enrich the atmosphere with oxygen, making it easier for air-breathing animals to colonize dry land. Plants also established early symbiotic relationships with fungi, creating mycorrhizae: a relationship in which the fungal network of filaments increases the efficiency of the plant root system, and the plants provide the fungi with byproducts of photosynthesis.

 

Career Connection

Paleobotanist: How organisms acquired traits that allow them to colonize new environments—and how the contemporary ecosystem is shaped—are fundamental questions of evolution. Paleobotany (the study of extinct plants) addresses these questions through the analysis of fossilized specimens retrieved from field studies, reconstituting the morphology of organisms that disappeared long ago. Paleobotanists trace the evolution of plants by following the modifications in plant morphology: shedding light on the connection between existing plants by identifying common ancestors that display the same traits. This field seeks to find transitional species that bridge gaps in the path to the development of modern organisms. Fossils are formed when organisms are trapped in sediments or environments where their shapes are preserved. Paleobotanists collect fossil specimens in the field and place them in the context of the geological sediments and other fossilized organisms surrounding them. The activity requires great care to preserve the integrity of the delicate fossils and the layers of rock in which they are found.

One of the most exciting recent developments in paleobotany is the use of analytical chemistry and molecular biology to study fossils. Preservation of molecular structures requires an environment free of oxygen, since oxidation and degradation of material through the activity of microorganisms depend on its presence. One example of the use of analytical chemistry and molecular biology is the identification of oleanane, a compound that deters pests. Up to this point, oleanane appeared to be unique to flowering plants; however, it has now been recovered from sediments dating from the Permian, much earlier than the current dates given for the appearance of the first flowering plants. Paleobotanists can also study fossil DNA, which can yield a large amount of information, by analyzing and comparing the DNA sequences of extinct plants with those of living and related organisms. Through this analysis, evolutionary relationships can be built for plant lineages.

Some paleobotanists are skeptical of the conclusions drawn from the analysis of molecular fossils. For example, the chemical materials of interest degrade rapidly when exposed to air during their initial isolation, as well as in further manipulations. There is always a high risk of contaminating the specimens with extraneous material, mostly from microorganisms. Nevertheless, as technology is refined, the analysis of DNA from fossilized plants will provide invaluable information on the evolution of plants and their adaptation to an ever-changing environment.

 

The Major Divisions of Land Plants

The green algae and land plants are grouped together into a subphylum called the Streptophyta, and thus are called Streptophytes. In a further division, land plants are classified into two major groups according to the absence or presence of vascular tissue, as detailed in Figure 25.6. Plants that lack vascular tissue, which is formed of specialized cells for the transport of water and nutrients, are referred to as non-vascular plants. Liverworts, mosses, and hornworts are seedless, non-vascular plants that likely appeared early in land plant evolution. Vascular plants developed a network of cells that conduct water and solutes. The first vascular plants appeared in the late Ordovician (500 to 435 MYA) and were probably similar to lycophytes, which include club mosses (not to be confused with the mosses) and the pterophytes (ferns, horsetails, and whisk ferns). Lycophytes and pterophytes are referred to as seedless vascular plants, because they do not produce seeds. The seed plants, or spermatophytes, form the largest group of all existing plants, and hence dominate the landscape. Seed plants include gymnosperms, most notably conifers, which produce “naked seeds,” and the most successful of all plants, the flowering plants (angiosperms). Angiosperms protect their seeds inside chambers at the center of a flower; the walls of the chamber later develop into a fruit.

 

Visual Connection

Figure 25.6 Streptophytes. This table shows the major divisions of green plants.

Which of the following statements about plant divisions is false?

1. Lycophytes and pterophytes are seedless vascular plants.
2. All vascular plants produce seeds.
3. All non-vascular embryophytes are bryophytes.
4. Seed plants include angiosperms and gymnosperms.

Evolution Connection

Building Phylogenetic Trees with Analysis of DNA Sequence Alignments: All living organisms display patterns of relationships derived from their evolutionary history. Phylogeny is the science that describes the relative connections between organisms, in terms of ancestral and descendant species. Phylogenetic trees, such as the plant evolutionary history shown in Figure 26.7, are tree-like branching diagrams that depict these relationships. Species are found at the tips of the branches. Each branching point, called a node, is the point at which a single taxonomic group (taxon), such as a species, separates into two or more species.

Figure 26.7 Plant phylogeny. This phylogenetic tree shows the evolutionary relationships of plants.

 

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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 thick 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 contribute to the success of terrestrial plants. The first adaptation is that the life cycle in all land plants exhibits the alternation of generations, a sporophyte in which the spores are formed and a gametophyte that produces gametes. Second is the antheridia and archegonia, which are the male and female reproductive structures (respectively) found on the gametophyte life stage. The third adaptation is the development of true tissues, including the apical meristem tissue in roots and shoots and parenchyma ground tissue. Fourth is the evolution of a waxy cuticle to resist desiccation (absent from some mosses). These adaptations all contribute to the success of the land plants, but are noticeably lacking in the closely related green algae—another reason for the debate over their placement in the plant kingdom. They are also not all found in the seedless plants, which can be regarded as representing an intermediate stage in adaptation to land.

 

Alternation of Generations

All sexually reproducing organisms have both haploid and diploid cells in their life cycles. In organisms with haplontic life cycles, the haploid stage is dominant, while in organisms with a diplontic life cycle, the diploid stage is the dominant life stage. Dominant in this context means both the stage in which the organism spends most of its time, and the stage in which most mitotic cell reproduction occurs—the multicellular stage. In haplontic life cycles, the only diploid cell is the zygote, which undergoes immediate meiosis to restore the haploid state. In diplontic life cycles, the only haploid cells are the gametes, which combine to restore the diploid state at their earliest convenience. Humans, for example, are diplontic. Alternation of generations describes a life cycle in which an organism has both haploid and diploid multicellular stages (Figure 25.2). This type of life cycle, which is found in all plants, is described as haplodiplontic.

Figure 25.2 Alternation of generations between the 1n gametophyte and 2n sporophyte is shown. Mitosis occurs in both gametophyte and sporophyte generations. Diploid sporophytes produce haploid spores by meiosis, while haploid gametophytes produce gametes by mitosis. (credit: Peter Coxhead)

In alternation of generations, the multicellular haploid form, known as a gametophyte, is followed in the developmental sequence by a multicellular diploid form, 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 seed plants. The evolution of the land plants is marked by increasing prominence of the sporophyte generation. The sporophyte stage is barely noticeable in non-vascular plants (the collective term for the plants that include the liverworts and mosses). In the seed plants, the sporophyte phase can be a towering tree, as in 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 sporophyte. This distinguishing feature of land plants gave the group its alternate name of embryophytes.

 

Sporophytes Produce Spores

The sporophyte of land plants is diploid and results from syngamy (fusion) of two gametes. The sporophyte bears the sporangia (singular, sporangium). The term “sporangia” literally means “a vessel for spores,” as it is a reproductive sac in which spores are formed (Figure 25.3). Inside the multicellular sporangia, the diploid sporocytes, or mother cells, produce haploid spores by meiosis, during which the 2n chromosome number is reduced to 1n (note that in many plants, chromosome number is complicated by polyploidy: 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, eventually generating a multicellular gametophyte by mitosis. The cycle then begins anew.

Plants that produce only one type of spore are called homosporous and the resultant gametophyte produces both male and female gametes, usually on the same individual. Non-vascular plants are homosporous, and the gametophyte is the dominant generation in the life cycle. Plants that produce two types of spores are called heterosporous. 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. A few seedless vascular plants and all seed plants are heterosporous, and the sporophyte is the dominant generation.

Figure 25.3 Sporangia. Spore-producing sacs called sporangia grow at the ends of long, thin stalks in this photo of the moss Esporangios bryum. (credit: Javier Martin)

 

Gametophytes Produce Gametes

Gametangia (singular, gametangium) are structures observed on multicellular haploid gametophytes. In the gametangia, precursor cells give rise to gametes by mitosis. The male gametangium, called the antheridium, releases sperm. Seedless plants produce sperm equipped with flagella that enable them to swim in a moist environment to the archegonium: the female gametangium. The embryo develops inside the archegonium as the sporophyte. In some plants, the antheridia and archegonia can be found on the same individual, while others have the gametangia separated. Gametangia are prominent in seedless plants, but have been lost in many seed plants. In seed plants, the gametophytes are microscopic and contained within pollen grains (male gametophytes) or ovules (female gametophyte). Some green algae also possess antheridia and archegonia, but it has been proposed that this is an example of homoplasy rather than homology due to the differences in morphology when compared to the archegonia and antehridia of land plants.

 

Plant Tissues

Another trait that distinguished land plants from green algae is the presence of true tissues. Plants are multicellular eukaryotes with tissue systems made of various cell types that carry out specific functions. Plant tissue systems fall into one of two general types: meristematic tissue, and permanent (or non-meristematic) tissue. Cells of the meristematic tissue are found in meristems, which are plant regions of continuous cell division and growth. In contrast, permanent tissue consists of plant cells that are generally no longer actively dividing.

Shoots and roots of plants increase in length through rapid cell division in a tissue called the apical meristem, which is a small mitotically active zone of cells found at the shoot tip or root tip (Figure 25.4). 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. 

Figure 25.4 Apical meristem at a root tip. 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.

 

Meristems produce cells that quickly differentiate, or specialize, and become permanent tissue. Such cells take on specific roles and often lose their ability to divide further. They differentiate into three main types: dermal, vascular, and ground tissue. Dermal tissue covers and protects the plant, and vascular tissue transports water, minerals, and sugars to different parts of the plant. Ground tissue serves as a site for photosynthesis, provides a supporting matrix for the vascular tissue, and helps to store water and sugars. Parenchyma is a type of ground tissue found in all land plants.

It is important to note that all these different tissues cannot be found in every land plant. For example, nonvascular plants (mosses, liverworts, and hornworts) do not have vascular tissue. That said, all land plants do show some degree of tissue differentiation, which is another trait that distinguished them from algae. While some green algae do have apical meristems, most algae do not have true tissues derived from the same embryological pathways as land plants. The tissue differentiation seen in land plants provided the physiological efficiency needed to specialize and adapt to the variety of niches on land.

 

Cuticle

In land plants, a waxy, waterproof cover called a cuticle protects the leaves and stems from desiccation. However, the cuticle also prevents intake of carbon dioxide needed for the synthesis of carbohydrates through photosynthesis. To overcome this, stomata or pores that open and close to regulate traffic of gases and water vapor appeared in plants as they moved away from moist environments into drier habitats.