25.1: Early Plant Life

25.1.1: Early Plant Life

Learning Objective

Describe the pervasiveness of seedless plants during the history of the kingdom Plantae

Key Points

• 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.

Key Terms

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.

25.1.2: Evolution of Land Plants

Learning Objective

Summarize the development of adaptations in land plants

Key Points

• 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.

Key Terms

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 .

The Rhynie chert sedimentary rock deposit

This Rhynie chert contains fossilized material from vascular plants. The area inside the circle contains bulbous underground stems called corms and root-like structures called rhizoids.

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 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 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.

25.1.3: Plant Adaptations to Life on Land

Learning Objective

Discuss how lack of water in the terrestrial environment led to significant adaptations in plants

Key Points

• While some plants remain dependent on a moist and humid environment, many have adapted to a more arid climate by developing tolerance or resistance to drought conditions.
• Alternation of generations describes a life cycle in which an organism has both haploid (1n) and diploid (2n) multicellular stages, although in different species the haploid or diploid stage can be dominant.
• The life on land presents significant challenges for plants, including the potential for desiccation, mutagenic radiation from the sun, and a lack of buoyancy from the water.

Key Terms

desiccation tolerance

the ability of an organism to withstand or endure extreme dryness, or drought-like condition

alternation of generation

the life cycle of plants with a multicellular sporophyte, which is diploid, that alternates with a multicellular gametophyte, which is haploid

Plant Adaptations to Life on Land

As organisms adapted to life on land, they had to contend with several challenges in the terrestrial environment. The cell's interior is mostly water: in this medium, 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 organisms exposed to air. Even when parts of a plant are close to a source of water, the aerial structures are prone to desiccation. 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 the ultraviolet rays of sunlight. Additionally, the male gametes must reach the female gametes using new strategies because swimming is no longer possible. As such, both gametes and zygotes must be protected from desiccation. Successful land plants have developed strategies to face all of these challenges. Not all adaptations appeared at once; some species never moved very far from the aquatic environment, although others went on to conquer the driest environments on Earth.

Despite these survival challenges, life on land does offer 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 water since it diffuses faster in air. Third, land plants evolved before land animals; therefore, until dry land was also 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 far from an abundant source of water and developed survival strategies to combat dryness. One of these strategies is called desiccation tolerance. Many mosses 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 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 and developed 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, which gives 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 (n represents the number of copies of chromosomes) . Haplontic refers to a lifecycle in which there is a dominant haploid stage (1n), while diplontic refers to a lifecycle in which the diploid (2n) 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. In fact, the sporophyte stage is barely noticeable in lower plants (the collective term for the plant groups of mosses, liverworts, and lichens). Alternatively, the gametophyte stage can occur in a microscopic structure, such as a pollen grain, in the higher plants (a common collective term for the vascular plants). Towering trees are the diplontic phase in the life cycles of plants such as sequoias and pines.

Alternation of generations of plants

Plants exhibit an alternation of generations between a 1n gametophyte and 2n sporophyte.

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.

25.1.4: Sporophytes and Gametophytes in Seedless Plants

Learning Objective

Describe the role of the sporophyte and gametophyte in plant reproduction

Key Points

• The diploid stage of a plant (2n), the sporophyte, bears a sporangium, an organ that produces spores during meiosis.
• Homosporous plants produce one type of spore which develops into a gametophyte (1n) with both male and female organs.
• Heterosporous plants produce separate male and female gametophytes, which produce sperm and eggs, respectively.
• In seedless plants, male gametangia (antheridium) release sperm, which can then swim to and fertilize an egg at the female gametangia (archegonia); this mode of reproduction is replaced by pollen production in seed plants.

Key Terms

syngamy

the fusion of two gametes to form a zygote

sporopollenin

a combination of biopolymers observed in the tough outer layer of the spore and pollen wall

gametangium

an organ or cell in which gametes are produced that is found in many multicellular protists, algae, fungi, and the gametophytes of plants

gametophyte

a plant (or the haploid phase in its life cycle) that produces gametes by mitosis in order to produce a zygote

sporophyte

a plant (or the diploid phase in its life cycle) that produces spores by meiosis in order to produce gametophytes

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": it is a reproductive sac that contains spores. 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.

Sporangia

Spore-producing sacs called sporangia grow at the ends of long, thin stalks in this photo of the moss Esporangios bryum.

Two different spore-forming methods are used 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 (1n) 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.

Lifecycle of heterosporous plants

Heterosporous plants produce two morphologically different types of spores: microspores, which develop into the male gametophyte, and megaspores, which develop into the female gametophyte.

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, also forms spores that contain sporopollenin.

Gametangia in Seedless Plants

Gametangia (singular, gametangium) are organs 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 replaced by pollen grains in seed-producing plants.

25.1.5: Structural Adaptations for Land in Seedless Plants

Learning Objective

Discuss the primary structural adaptations made by plants to living on land

Key Points

• Many plants developed a vascular system: to distribute water from the roots (via the xylem) and sugars from the shoots (via the phloem) throughout the entire plant.
• An apical meristem enables elongation of the shoots and roots, allowing a plant to access additional space and resources.
• Because of the waxy cuticle covering leaves to prevent water loss, plants evolved stomata, or pores on the leaves, which open and close to regulate traffic of gases and water vapor.
• Plants evolved pathways for the synthesis of complex organic molecules, called secondary metabolites, for protection from both UV lights and predators.

Key Terms

meristem

the plant tissue composed of totipotent cells that allows plant growth

stoma

a pore found in the leaf and stem epidermis used for gaseous exchange

xylem

a vascular tissue in land plants primarily responsible for the distribution of water and minerals taken up by the roots; also the primary component of wood

phloem

a vascular tissue in land plants primarily responsible for the distribution of sugars and nutrients manufactured in the shoot

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).

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 . 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.

Apical meristem

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.

Vascular structures

In small plants such as single-celled algae, simple diffusion suffices to distribute water and nutrients throughout the organism. However, for plants to develop larger forms, the evolution of vascular tissue for the distribution of water and solutes was a prerequisite. The vascular system contains xylem and phloem tissues. Xylem conducts water and minerals absorbed from the soil up to the shoot, while phloem transports food derived from photosynthesis throughout the entire plant. A root system evolved to take up water and minerals from the soil, while anchoring the increasingly taller shoot in the soil.

Additional land plant adaptations

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. This presented an additional challenge to land colonization, which was met by the evolution of biosynthetic pathways for the synthesis of protective flavonoids and other compounds: 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.

25.1.6: The Major Divisions of Land Plants

Learning Objective

Identify the major divisions of land plants

Key Points

• Non-vascular plants, or bryophytes, appeared early in plant evolution and reproduce without seeds; they include mosses, liverworts, and hornworts.
• Vascular plants are subdivided into two classes: seedless plants, which probably evolved first (including lycophytes and pterophytes), and seed plants.
• Seed-producing plants include gymnosperms, which produce "naked" seeds, and angiosperms, which reproduce by flowering.

Key Terms

spermatophyte

any plant that bears seeds rather than spores

embryophyte

any member of the subkingdom Embryophyta; most land plants

bryophyte

seedless, nonvascular plants that are the closest extant relative of early terrestrial plants

The Major Divisions of Land Plants

The green algae, known as the charophytes, and land plants are grouped together into a subphylum called the Streptophytina and are, therefore, called Streptophytes. Land plants, which are called embryophytes, are classified into two major groups according to the absence or presence of vascular tissue . 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 or bryophytes. Non-vascular embryophytes probably appeared early in land plant evolution and are all seedless. These plants include liverworts, mosses, and hornworts.

Major divisions of land plants

Land plants are categorized by presence or absence of vascular tissue and their reproduction with or without the use of seeds.

In contrast, vascular plants developed a network of cells, called xylem and phloem, that conduct water and solutes throughout the plant. The first vascular plants appeared in the late Ordovician period of the Paleozoic Era (approximately 440-485 million years ago). These early plants were probably most similar to modern day lycophytes, which include club mosses (not to be confused with the mosses), and pterophytes, which include ferns, horsetails, and whisk ferns. Lycophytes and pterophytes are both referred to as seedless vascular plants because they do not produce any seeds.

The seed producing plants, or spermatophytes, form the largest group of all existing plants, dominating the landscape. Seed-producing plants include gymnosperms, most notably conifers, which produce "naked seeds," and the most successful of all modern-day plants, angiosperms, which are the flowering plants. Angiosperms protect their seeds inside chambers at the center of a flower; the walls of the chamber later develop into a fruit.

Attributions

• Early Plant Life
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  • "OpenStax College, Biology. October 17, 2013." http://cnx.org/content/m44633/latest/?collection=col11448/latest. OpenStax CNX CC BY 3.0.
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  • "OpenStax College, Introduction. October 17, 2013." http://cnx.org/content/m44633/latest/Figure_25_00_01.jpg. OpenStax CNX CC BY 3.0.
• Evolution of Land Plants
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• Plant Adaptations to Life on Land
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• Sporophytes and Gametophytes in Seedless Plants
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• Structural Adaptations for Land in Seedless Plants
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  • "OpenStax College, Early Plant Life. November 17, 2013." http://cnx.org/content/m44635/latest/Figure_25_01_03.jpg. OpenStax CNX CC BY 3.0.
• The Major Divisions of Land Plants
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  • "spermatophyte." http://en.wiktionary.org/wiki/spermatophyte. Wiktionary CC BY-SA 3.0.
  • "OpenStax College, Early Plant Life. October 17, 2013." http://cnx.org/content/m44635/latest/Figure_25_01_05.jpg. OpenStax CNX CC BY 3.0.