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The geologic periods of the Paleozoic are marked by changes in the plant life that inhabited the earth.
- Summarize the development of adaptations in land plants
- Land plants first appeared during the Ordovician period, more than 500 million years ago.
- The evolution of plants occurred by a stepwise development of physical structures and reproductive mechanisms such as vascular tissue, seed production, and flowering.
- Paleobotonists trace the evolution of plant morphology through a study of the fossil record in the context of the surrounding geological sediments.
- Paleobotany: the branch of paleontology or paleobiology dealing with the recovery and identification of plant remains from geological contexts
- mycorrhiza: a symbiotic association between a fungus and the roots of a vascular plant
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 evolution of plants occurred by a gradual development of novel structures and reproduction mechanisms. Embryo protection developed prior to the development of vascular plants which, in turn, evolved before seed plants and flowering plants. 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, where embedded fossils of some of the earliest vascular plants have been identified.
Gradual evolution of land plants: The adaptation of plants to life on land occurred gradually through the stepwise development of physical structures and reproduction mechanisms
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, which sheds 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. Paleobotanists collect fossil specimens in the field and place them in the context of the geological sediments and other fossilized organisms surrounding them.
Paleobotanists distinguish between extinct species, as fossils, and extant species, which are still living. The extinct vascular plants, classified as zosterophylls and trimerophytes, most probably lacked true leaves and roots, forming low vegetation mats similar in size to modern-day mosses, although some trimetophytes 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 Cooksoniapossessed 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 in 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. The plants provide the fungi with byproducts of photosynthesis.
25.1B: Evolution of Land Plants - Biology
A diverse array of seedless plants still populate and thrive in the world today, particularly in moist environments.
Describe the pervasiveness of seedless plants during the history of the kingdom Plantae
- Non-vascular seedless plants, or bryophytes, are the group of plants that are the closest extant relative of early terrestrial plants.
- The vast majority of terrestrial plants today are seed plants, which tend to be better adapted to the arid land environment.
- Seedless plants are classified into three main catagories: green algae, seedless non- vascular plants, and seedless vascular plants.
- vascular plant: any plant possessing vascular tissue (xylem and phloem), including ferns, conifers, and flowering plants
- bryophyte: seedless, nonvascular plants that are the closest extant relative of early terrestrial plants
Introduction to Early Plant Life
An incredible variety of seedless plants populates the terrestrial landscape. Mosses may grow on a tree trunk and horsetails may display their jointed stems and spindly leaves across the forest floor. Today, however, seedless plants represent only a small fraction of the plants in our environment. The kingdom Plantae constitutes a large and varied group of organisms with more than 300,000 species of cataloged plants. Of these, more than 260,000 are seed plants. However, three hundred million years ago, seedless plants dominated the landscape and grew in the enormous swampy forests of the Carboniferous period. Their decomposition created large deposits of coal that we mine today.
Horsetails are seedless plants: Seedless plants, like these horsetails (Equisetum sp.), thrive in damp, shaded environments under a tree canopy where dryness is rare.
Current evolutionary thought holds that all plants, green algae as well as land dwellers, are monophyletic that is, they are descendants of a single common ancestor. The evolutionary transition from water to land imposed severe constraints on plants. They had to develop strategies: to avoid drying out, to disperse reproductive cells in air, for structural support, and for capturing and filtering sunlight. While seed plants developed adaptations that allowed them to populate even the most arid habitats on Earth, full independence from water did not happen in all plants. Most seedless plants still require a moist environment.
Seedless plants are classified into three main categories: green algae, seedless non-vascular plants, and seedless vascular plants. Seedless non-vascular plants (bryophytes), such as mosses, are the group of plants that are the closest extant relative of early terrestrial plants. Seedless vascular plants include horsetails and ferns.
25.1 Early Plant Life
By the end of this section, you will be able to do the following:
- 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.
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.
Link to Learning
Go to this article to get a more in-depth view of the Charophytes.
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 an apical meristem tissue in roots and shoots. Third is the evolution of a waxy cuticle to resist desiccation (absent from some mosses). Finally cell walls with lignin to support structures off the ground. 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 mosses, 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 .
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 .
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). 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. 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.
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.
The spores of seedless plants are surrounded by thick cell walls containing a tough polymer known as sporopollenin . As the name suggests, it is also found in the walls of pollen grains. 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, in which pollen is the male gametophyte, 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 charophyte Coleochaetes also 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. 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. Gametangia are prominent in seedless plants, but are absent or rudimentary in seed plants.
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.
Additional Land Plant Adaptations
As plants adapted to dry land and became independent from the constant presence of water in damp habitats, new organs and structures made their appearance. Early land plants did not grow more than a few inches off the ground, competing for light on these low mats. By developing a shoot and growing taller, individual plants captured more light. Because air offers substantially less support than water, land plants incorporated more rigid molecules in their stems (and later, tree trunks). In small plants such as single-celled algae, simple diffusion suffices to distribute water and nutrients throughout the organism. However, for plants to evolve larger forms, the evolution of a conductive tissue for the distribution of water and solutes was a prerequisite. The evolution of vascular tissue in plants met both of these needs. The vascular system contains two types of conductive tissue: xylem and phloem. Xylem conducts water and minerals absorbed from the soil up to the shoot, while phloem transports food derived from photosynthesis throughout the entire plant. In xylem, the cells walls are reinforced with lignin, whose tough hydrophobic polymers help prevent the seepage of water across the xylem cell walls. Lignin also adds to the strength of these tissues in supporting the plant. The vascular tissues extend into the root of land plants. The root system evolved to take up water and minerals from the soil, and to anchor the increasingly taller shoot in the soil.
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.
Water filters ultraviolet-B (UVB) light, which is harmful to all organisms, especially those that must absorb light to survive. This filtering does not occur for land plants. Exposure to damaging radiation presented an additional challenge to land colonization, which was met by the evolution of biosynthetic pathways for the synthesis of protective flavonoids and other pigments that absorb UV wavelengths of light and protect the aerial parts of plants from photodynamic damage.
Plants cannot avoid being eaten by animals. Instead, they synthesize a large range of poisonous secondary metabolites: complex organic molecules such as alkaloids, whose noxious smells and unpleasant taste deter animals. These toxic compounds can also cause severe diseases and even death, thus discouraging predation. Humans have used many of these compounds for centuries as drugs, medications, or spices. In contrast, as plants co-evolved with animals, the development of sweet and nutritious metabolites lured animals into providing valuable assistance in dispersing pollen grains, fruit, or seeds. Plants have been enlisting animals to be their helpers in this way for hundreds of millions of years.
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.
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.
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.
Adaptations to alternation of generations in land plants
As we’ve previously discussed, all eukaryote life cycles include a haploid stage and a diploid stage. Usually one of these stages is large and multicellular (the organism we can see by eye), while the other is small and unicellular. All land plants (and *some* green algae) reproduce via the alternation of generations life cycle, where both the haploid and the diploid stage of an organism are multicellular: the haploid multicellular form, known as a gametophyte, is followed in the life cycle 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 v ascular plants. The sporophyte stage is barely noticeable in nonvascular plants . Towering trees are the diplontic phase in the lifecycles of plants such as sequoias and pines. The image below shows a simplified version of the alternation of generations life cycle:
Alternation of Generations. Image credit: Menchi, Wikimedia Commons. https://en.wikipedia.org/wiki/File:Sporic_meiosis.png
Though all plants display an alternation of generations life cycle, there are significant variations in different lineages of plants, consistent with their evolutionary history and order of origination:
- In seedless non-vascular plants, or bryophytes, the haploid gametophyte is larger than the sporophyte (the plant structure that you see is the gametophyte) this is a gametophyte-dominated life cycle. (By “dominated” we mean “the stage of the plant you can see by eye.”) The sporophyte is attached to and dependent on the gametophyte for water and nutrients.
- In seedless vascular plants (such as ferns), the sporophyte is larger than the gametophyte (the plant structure that you see is the sporophyte), but the gametophyte is free-living and independent from the diploid sporophyte.
- The life cycle of gymnosperms (conifers) and angiosperms (flowering plants) is dominated by the sporophyte stage (the plant structure that you see is the sporophyte), with the gametophyte remaining attached to and dependent on the sporophyte (reverse of bryophytes).
- Though they both have sporophyte-dominated life cycles, angiosperms and gymnosperms differ in that angiosperms have flowers, fruit-covered seeds, and double fertilization, while gymnosperms do not have flowers, have “naked” seeds, and do not have double fertilization.
The video below describes the features of nonvascular plants (mosses, liverworts, hornworts), and their alternation of generations life cycle:
The video below describes the features of vascular plants and their alternation of generations life cycle:
Alternation of Generations Life Cycle
To form a sporophyte, two haploid gametes come together to form a diploid zygote. Typically, haploid organisms are defined by having an “n” number of chromosomes. When two gametes of the same species come together, they each have n chromosomes. Therefore, the diploid zygote which forms is considered to have 2n worth of genetic material, or exactly twice as much. Not only is there twice as much DNA, but it represents codes for the same proteins in the same organism. The sporophyte is a multicellular organism formed from multiple rounds of mitosis on the zygote. Thus, the sporophyte individual remains a 2n organism.
Then, when the sporophyte reaches maturity, a key point in the alternation of generations takes place. The sporophyte develops organs, known as sporangia. These specialized reproductive organs are used to create single-celled, haploid spores. These cells will be released into the air or water and carried away. When they reach a suitable environment, they will begin the process of developing into the gametophyte.
This represents the next generation in the alternation of generations, as the haploid spore is created. The spore is technically a new organism, and has only half the DNA as the parent organism. This spore will undergo successive rounds of mitosis to form a new multicellular individual, the gametophyte. Where the sporophyte generation creates spores, the gametophyte generation creates gametes. Gametes are produced by special organs on the gametophyte, the gametangia. These gametes are then broadcast into the environment, or transferred between plants.
When they find an opposite gamete, they begin the process of fusing to form another zygote. This zygote will eventually become a sporophyte, and the alternation of generations will keep turning. While this is a simplistic version of the alternation of generations, there are many complexities, which will be discussed below. Because of these complexities, and because all plants undergo some version of alternation of generations, scientists prefer to refer to other aspects of their reproductive cycles to define the species.
I. The singularity of plant terrestrialization
Given that plants evolved from within streptophytes, there is considerable interest in trying to reconstruct early land plant evolution through the lens of streptophyte algae. Among the most fundamental questions in the field are: which specific lineage of streptophyte algae gave rise to the land plants and what was it about the biology of this lineage that enabled its conquest of land? The answer to the first question is now in hand. The paraphyletic streptophyte algae can be divided into the lower-branching KCM-grade (Klebsormidiophyceae, Chlorokybophyceae and Mesostigmatophyceae) and the higher-branching ZCC-grade (Zygnematophyceae, Coleochaetophyceae, and Charophyceae see Box 1, Fig. 1 also see de Vries et al., 2016 ). For some time it seemed as though there had been a stepwise increase in organismal complexity, from a unicellular alga like extant Mesostigma (see Marin & Melkonian, 1999 ), through to simple filamentous forms such as Klebsormidium and Zygnema, on to pseudoparenchymatous apical cell-bearing Coleochaete, and, finally, to a complex multicellular organism like Chara, a stonewort with rhizoids and stem-like structures (Fig. 1). However, recent phylogenomic analyses show a sister relationship between land plants and the Zygnematophycae, an enigmatic group of unicellular and filamentous algae that live in a variety of freshwater and terrestrial environments (Wodniok et al., 2011 Timme et al., 2012 Ruhfel et al., 2014 Wickett et al., 2014 ). Given that the Zygnematophyceae have the least complex bodyplan of all higher-branching streptophyte algae (i.e. the ZCC grade), this comes as something of a surprise. As we shall see, inferring the evolution of traits along the trajectory of streptophyte evolution has proved challenging.
Box 1. Glossary
Chloroplastida – the monophyletic ‘green’ lineage consisting of the chlorophytes (green algae) and streptophytes.
Homoiohydry – a dehydration-avoidance mechanism based on active maintenance of water homeostasis – especially under water-limiting conditions the predominant mechanism employed by vascular land plants to tolerate water limitations.
KCM grade – a paraphylum consisting of the basal-branching streptophyte algal clades Klebsormidiophyceae, Chlorokybophyceae, and Mesostigmatophyceae.
Phragmoplast – a scaffold of cytoskeletal components and vesicles that is formed during anaphase/telophase a textbook characteristic of the land plant-type cell division.
Poikilohydry – passive equilibration of the cell's water status to that of the environment can lead to desiccation under water-limiting conditions, which is tolerated by only some photosynthetic eukaryotes such as certain algae (including some streptophyte algae), lichens, mosses and very few vascular plants.
Preprophase band – an actin and tubulin scaffold that forms during the end of the G2 phase of Zygnematophyceae and land plants (for a comprehensive discussion, see Buschmann & Zachgo, 2016 ).
Primary plasmodesmata – cell wall channels that are formed during cell division (as opposed to secondary plasmodesmata, which are formed independently of cell division).
Sporopollenin – the highly robust material that coats spores of algae, embryophytes, and fungi. It consists of a complex mixture of biopolymers, including (but not limited to) phenolic compounds and fatty acids best known as the robust coating of pollen.
Streptophytes – a monophyletic group uniting the streptophyte algae (also known as charophytes) and land plants.
ZCC grade – a paraphylum of the higher-branching streptophyte algal clades Zygnematophyceae, Coleochaetophyceae, and Charophyceae, which are sister to land plants.
The second question is much more difficult. Evolution is a dynamic process extant organisms are snapshots in time and provide only static, and often misleading, information about how they came to be. In order to infer the features of streptophyte algae that were relevant to the conquest of land, one needs not only to consider present-day Zygnematophycae, but also to look at their ancestry. To identify informative features, it is crucial to consider what plant terrestrialization actually means. In thinking about the success of land plants, obviously adaptive traits such as the presence of desiccation-tolerant seeds, vascular tissues (which conduct water and provide stability), and gas-exchanging stomata quickly come to mind. These are certainly essential features of the (homoiohydric see Box 1) plant cormus – the true plant body – and > 400 million-yr-old fossils exhibiting many of these properties have been found (reviewed by Gensel, 2008 ). However, the cormus arose within land plants the aforementioned features are hence the subject of the evolution of plants on land. Towards a complete understanding of land plant evolution we must also consider what came about before such features arose. The question is more than what the first vegetative body looked like. It is a physiological question about how the organisms that eventually gave rise to land plants overcame the challenges associated with dry land (reviewed by Delwiche & Cooper, 2015 ). Cellular physiology and bodyplan evolution are in fact two sides of the same coin: a more complex bodyplan allows for cellular differentiation, which later proved to be essential for the evolution of specific tissues (e.g. water-conducing vasculature) and land plant homoiohydry. But the algal ancestor of embryophytes was a poikilohydric organism – just like any other (extant) terrestrial microalga, which can also grow in diverse and challenging environments.
The algal progenitor of embryophytes did more than just dwell in a terrestrial habitat such as moist soil or rock. It conquered all parts terra firma, presumably in a stepwise fashion. What physiological properties allowed the first land plants to thrive such that the complex adaptations so familiar to plant biologists could evolve? Here we discuss some of the concepts and challenges associated with inferring early steps in land plant evolution.
What Your Textbook Says about the Origin of Plants
|Plants appear as fossils over 400 million years ago.||458, 560||420||551||265, 490, 502||7:1, 7:3|
|Plants evolved from green algae and invaded the vacant land niche.||559, 666||420, T420||430, 511, T512, T514, 553, 554, 555||265, 422, 502||7:2, 7:3, 7:4|
|Many adaptations were needed as plants moved from water onto land.||562, 564, 580||418, T418, 420–425||430, T549, 554, 555, 643, 644, T645||502– 505||7:4|
|Many symbiotic relationships evolved to support plants on land.||—||411||437–438, T438, 542, T553, 615||T266, 362||3:14|
|Early forests were dominated by giant ferns and cycads that formed fossil fuels. These were later replaced by seed plants.||566, 567, 584||427||T77, 144, 430, 431, 561, 566, 567||266||7:3|
|The first fossils of vascular plants occur at 375 million years ago.||586–587||427||554, 560, T560, 562, 566||510||7:3|
|The first fossil seed plants occur at 360 million years ago, as seed plants replace ferns and cycads.||596–597||431-433||564, 566, 568, T571||504||7:3|
|Flowering plants evolve slowly and appear in the fossils 130 million years ago.||569, 656||434, 436–437, T443, 445, 448, 470, 481–485||432, T432, 569, T612, 618||505, 514||7:3|
|Bristlecone pines in the Rocky Mountains are the oldest known organisms at 4,700 years old.||—||T454||T567||513, T570||7:5|
Note: Page numbers preceded by “T” indicate items from the teacher notes found in the margins of the Teacher’s Edition.