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18 EVOLUTION OF PLANTS EXTENDED LECTURE OUTLINE Plants (p. 366) 18.1 18.2 Adapting to Terrestrial Living (p. 366; Figs. 18.1, 18.2, 18.3, 18.4) A. Plants are multicellular, terrestrial autotrophs that occur on land and derive energy through photosynthesis. B. The green algae gave rise to the diverse land plants, which have now evolved into at least 266,000 different species. C. Three major challenges had to be overcome for living on land: organisms had to absorb minerals, conserve water, and develop a new mode of reproduction suitable for land. D. Absorbing Minerals 1. Many of the early plants had symbiotic fungi, such as the present-day mycorrhizae, to aid in the uptake of minerals and nutrients from rocky shores devoid of soil. E. Conserving Water 1. A waxy cuticle evolved to help retard water loss. 2. Gases could be exchanged with the atmosphere through tiny openings called stomata that could also be closed to further reduce water loss. F. Reproducing on Land 1. The development of spores was an evolutionary trend that protected reproductive cells from drying. 2. Transferring gametes from one plant to the next during sexual reproduction posed a problem that has been answered by the development of spores in many plant lines. 3. Haploid spores are produced by sporophytes, the diploid generation that alternates with the haploid gamete-producing gametophyte generation. 4. This alternation of generations solved many problems for plants reproducing sexually on land. Evolution of a Vascular System (p. 368; Fig. 18.5; Table 16.1) A. As plants evolved greater specializations on land, below-ground root systems developed, along with vascular tissue to connect above- and below-ground portions of plants. B. These “plumbing” systems are now specialized vascular systems that run from the very bottom of the roots to the tips of the shoots. C. Plants that have evolved vascular systems are the most common types of plants found on earth at the present. Seedless Plants (p. 370) 18.3 Nonvascular Plants (p. 370; Fig. 18.6) A. Liverworts and Hornworts 1. Some plants that never evolved vascular systems can still be found. 2. The liverworts and hornworts are two groups of primitive plants that completely lack any kind of vascular system. 3. These small plants can only survive in moist habitats. B. Primitive Conducting Systems: Mosses 1. Mosses, a third type of fairly primitive plant, managed to evolve a simple, soft-sided tube capable of conducting water over short distances. 2. Strengthened tubes evolved later as vascular tissue in the true vascular plants. 3. In the liverworts, hornworts, and mosses, the main plant body is the gametophyte, with the sporophyte being borne on the gametophyte. 94 18.4 18.5 The Evolution of Vascular Tissue (p. 371; Figs. 18.7, 18.8) A. Vascular plants have specialized cells that can conduct materials and fluids. B. The remaining nine phyla of plants have such vascular tissue, which enables them to grow to much greater size than the nonvascular plants. C. The fossil record indicates that vascular plants have been around for 430 million years and are different from nonvascular plants in that they spend most of their time in the sporophyte stage, have specialized conducting tissue, and have a specialized body form made up of roots, stems, and leaves. D. Two types of specialized conducting tissue can be found in vascular plants. 1. Phloem, made up of soft-walled sieve elements, conducts carbohydrates manufactured during photosynthesis from the leaves to the rest of the plant, and xylem, made up of hard-walled tracheary elements, conducts water from the roots upward to the rest of the plant. E. How vascular plants grow has changed somewhat over the course of evolution. 1. Primitive vascular plants exhibited primary growth only, which involves elongation of roots and stems from a growing point called the apical meristem. 2. Vascular plants of today still exhibit this type of growth, but are also able to grow in width, called secondary growth, as a result of a cylinder of meristem lying beneath the bark. 3. Secondary growth produces wood. Seedless Vascular Plants (p. 372; Figs. 18.9, 18.10) A. Four phyla of modern vascular plants lack seeds and rely on the presence of water to enable sperm to swim to their destination for fertilization. B. Ferns are among the most abundant of the seedless vascular plants, and are especially numerous in the tropics. C. The Life of a Fern 1. They exist as sporophytes with large leaves called fronds. 2. Spores are produced on the fronds and fall to the earth to develop into gametophytes. 3. On the small heart-shaped gametophytes, gametes are produced, fertilization occurs, and a new sporophyte grows. The Advent of Seeds (p. 374) 18.6 Evolution of Seed Plants (p. 374; Figs. 18.11, 18.12, 18.13) A. The evolution of seeds has greatly enhanced the ability of plants to survive on land. B. A seed protects the developing embryo when it is most vulnerable to drying out or predation. C. In seed plants, the gametophyte generation is even smaller than in seedless vascular plants, and gametophytes exist entirely within the sporophyte plant. 1. Microgametophytes, or pollen grains, are male gametophytes, and are transported by various means, depending upon the particular adaptations of a plant. 2. The female gametophyte is the megagametophyte, or ovule, which is fertilized only after pollination. D. The first seed plants were the gymnosperms with incompletely enclosed ovules. E. Angiosperms evolved later, with their ovules completely enclosed within a vessel called a carpel. F. The Structure of a Seed 1. A seed is made up of the embryo of the sporophyte surrounded by a hard, protective seed coat. 2. Nourishing endosperm is also housed within the seed coat to provide a food source until germination. 3. Seeds are adapted to life on land because they enable better dispersal, or colonization of new areas or better habitats, and because they protect the embryo during a period of dormancy when the embryo is waiting for better conditions before germination. 95 4. 18.7 Seeds are also tuned into the appropriate environmental cues that indicate when moisture and temperature are optimal before germination and growth of the embryo out of the seed. Gymnosperms (p. 376; Figs. 18.14, 18.15, 18.16) A. Gymnosperms are represented on earth today by four different phyla: conifers, cycads, ginkgos, and the gnetophytes. B. We are most familiar with the conifers whose needlelike leaves help resist water loss for these among the oldest and largest trees on earth. C. The Life of a Gymnosperm 1. Conifers form two kinds of cones: seed cones that bear female gametophytes and pollen cones that produce male gametophytes. 2. Conifers rely on the wind to transfer pollen to the seed cone. 3. Once the sperm cell inside the pollen fuses with the egg cell, a new zygote is formed that can eventually develop into a sporophyte. The Evolution of Flowers (p. 378) 18.8 18.9 18.10 18.11 Rise of the Angiosperms (p. 377; Fig. 18.17) A. Angiosperms are probably the most successful land plants, comprising 235,000 species or 90% of all land plants. B. Angiosperms evolved unique ways of transferring pollen for fertilization, which has ensured their survival, and special structures, called flowers, to attract and reward animal pollinators. C. The Flower 1. Flowers are structured into four concentric whorls. 2. The outermost whorl, made up of sepals, protects the flower while it is still a bud. 3. The second whorl is made up of the colorful petals that attract pollinators. 4. The third whorl is the male portion of the flower, made up of stamens that bear a pollenproducing swollen tip called the anther. 5. The innermost whorl is female in nature, and contains the carpel that houses ovules or egg cells, a long stalk called the style leading to the ovules, and a stigma at the tip that receives pollen. 6. When pollen arrives at the stigma, a pollen tube grows down through the style to reach the ovule. Why Are There Different Kinds of Flowers (p. 379; Fig. 18.18, 18.19) A. Some flower adaptations that have coevolved to attract and satisfy the needs of the pollinator include unique petal arrangements and colors, patterns of color, odor, and nectar rewards. B. Angiosperms that rely on wind pollination, like grasses, oaks, and birches, do not have large, showy, fragrant flowers. Improving Seeds: Double Fertilization (p. 380; Figs. 18.20, 18.21) A. Angiosperm seeds have a greatly improved food source for the growing embryo. B. Endosperm is a triploid nutritive source for the growing embryo, and forms as a result of double fertilization. C. One sperm nucleus fertilizes the ovule while another fuses with the polar nuclei to form the triploid endosperm. D. Endosperm divides very rapidly to ensure an adequate food supply for the embryo. E. In some angiosperms, food is stored in swollen, fleshy leaves called cotyledons. 1. Dicots are plants with two cotyledons, like oak and maple trees. 2. Monocots have only one cotyledon and other characteristics different from dicots. 3. Grasses are among the most successful monocots. Improving Seed Dispersal: Fruits (p. 382; Fig. 18.22) A. Fruits are a means of dispersing seeds. B. A fruit is a ripened, fleshy ovary containing the seeds. C. Animals eat fruit and pass the seeds in feces. D. Many specialized types of fruit can be found among the angiosperms. 96 LEARNING OBJECTIVES Recognize which group of algae gave rise to land plants and the nature of the problems this group had to overcome to live on land. List the adaptations that evolved in groups of land plants to conserve water and to enable reproduction on land. Describe liverworts, hornworts, and mosses. Explain the nature of vascular systems. Describe the life cycle of ferns. Discuss the advantages and adaptations of seeds. Compare gymnosperms with angiosperms. List the major groups of gymnosperms. Diagram the gymnosperm life cycle. Describe the structure of a flower. Discuss how angiosperms coevolved with their pollinators. Understand what is meant by “double fertilization.” Compare the structures of monocots with those of dicots. Explain the adaptive significance of fruits. KEY TERMS cuticle (p. 366) The cuticle is a waxy, waterproof substance that prevents water loss to the air. stomata (p. 366) In leaves and in some green stems, carbon dioxide enters through these openings that can be closed to prevent water loss. alternation of generations (p. 367) sporophyte (p. 367) Diploid generation; forms haploid spores by meiosis. gametophyte (p. 367) Haploid generation; forms haploid gametes by mitosis. liverwort (p. 370) hornwort (p. 370) moss (p. 370) People often confuse mosses with algae, calling growth on stream rocks “moss” when it is algae. vascular tissue (p. 371) primary growth (p. 371) This involves cell division at the tips of the shoots and roots. secondary growth (p. 371) Secondary growth is growth in girth. fern (p. 372) One of the four phyla of seedless vascular plants with 12,000 living species. seed (p. 374) A seed is an embryo with a watertight cover. gymnosperm (p. 376) conifers (p. 376) Conifers are one type of gymnosperm that produces its seeds in cones. cone (p. 376) angiosperm (p. 378) Angiosperms have an ovule that is completely enclosed by the sporophyte when fertilized. flower (p. 378) Flowers are the reproductive organs of angiosperms. endosperm (p. 380) Nutritious triploid tissue within the seed. double fertilization (p. 381) Fertilization of the zygote and of the endosperm is double fertilization. dicot (p. 381) monocot (p. 381) 97 LECTURE SUGGESTIONS AND ENRICHMENT TIPS 1. Seedless Vascular Plants. Few people give much notice to the seedless vascular plants, partly because they are small, and partly because of lack of knowledge about them. Do a “show and tell” demonstration of the seedless vascular plants, allowing students to examine them individually. Show them examples of Marchantia with its archegoniophores and antheriodiophores, and Mnium or similar moss species with its archegonial heads. Demonstration specimens of a variety of ferns and prepared slides of fern prothalli are useful in illustrating the stages of a fern life cycle. 2. Endangered Plants. Students are fairly well aware of the idea of endangered species, although many people automatically think of mammals and birds when they hear that phrase. What we don't often realize is the importance of endangered plants, and the effect of plant species extinctions on ecosystems. The fossil record indicates that five or six large bouts of extinction have occurred over the past 500 million years, and most of the organisms going extinct at once were animals. We tend to notice when animals in our vicinity become less numerous, but we are less apt to notice, or perhaps care about, the plants. However, plants are more important from an ecological standpoint because most insect and other animal species depend on plants in one way or another to stay alive. Probably 10% of the world's plant species are now facing extinction, and in a few short years, as many as 25% of the world's plant species can be expected to be threatened with extinction. Questions to consider as you discuss this with your students are: What can be done to slow down extinction rates of plants? Where are many extinctions occurring? (In the tropics, but examples can be found everywhere.) What will happen to the animals when the plants on which they depend become extinct? What can each of us do to help solve this problem? 3. Plants During the Time of the Dinosaur. What types of plants dominated the landscape during the day of the dinosaur? Illustrate your presentation with drawings of the types of plants present during various geologic periods. Bring in fossil specimens, or photographs of same, as available. During the Silurian period of the Paleozoic era (around 410 million years ago), the low-lying primitive vascular land plants first appeared. Seed ferns evolved during the Devonian period, approximately 390 million years ago. During the Carboniferous period, 286–360 million years ago, the great coal-forming forests dominated the earth. Ferns, horsetails, and club mosses were at their most numerous. Conifers appeared in the fossil record next, some 246–286 million years ago during the Permian period. At the beginning of the Mezozoic era, during the dawn of the dinosaurs, forests of ferns and gymnosperms covered the landscape, and cycads and ginkgoes first appeared in the fossil record. When dinosaurs reached immense sizes and numbers in the Jurassic period 150–200 million years ago, cycads and other gymnosperms flourished. From 70 to 150 million years ago, the flowering plants appeared, placental mammals evolved, and dinosaurs went extinct in massive numbers at the close of the Cretaceous period. From then on, the angiosperms and mammals have been the dominant organisms, along with modern insect groups and, most recently, humans. CHANGES TO THE NEW EDITION Refer to the Johnson instructor web site at http://www.mhhe.com/biosci/genbio/tlw4 for a complete list of changes to this edition. CRITICAL THINKING QUESTIONS 1. 2. 3. Describe the evidence that suggests land plants evolved from green algae. Explain why the evolution of the seed was a critical step in the domination of land by plants. In what geographical areas are gymnosperms, rather than angiosperms, dominant and why? 98 FILMS/MEDIA SUGGESTIONS (Telephone and fax numbers and/or web sites for the sources of the following materials are listed in the Appendix.) How Plants Are Classified. The history of plant classification and the current scheme for classifying plants are reviewed, with examples of each level given. 18 minutes. CLEARVUE/eav, WWCL330-1CV Simple Photosynthetic Organisms: From Algae to Ferns. The characteristics of the algae and the simplest land plants, mosses, liverworts, and ferns, as well as their evolution and adaptations are explored in this video. 18 minutes. CLEARVUE/eav, WWCL330-2CV Seed Plants: Gymnosperms and Angiosperms. How these plants differ structurally from simpler plants, and how they have evolved to live more successfully on land are the main topics presented in this video. How flowering plants differ from nonflowering plants is also discussed. 18 minutes. CLEARVUE/eav, WWCL330-3CV The above three videos are available as a group, The Development of Plants. CLEARVUE/eav, WWCL330-CV Angiosperms. Highlights of this video include the importance of angiosperms, the differences between monocots and dicots, annuals versus biennials and perennials, and animated sequences showing the life cycle of a flowering plant, pollen and egg formation, and double fertilization. 31 minutes. CLEARVUE/eav, WW5VH 1615 Gymnosperms. The first seed plants, the ginkgoes, cycads, conifers, and gnetophytes, are highlighted along with pollination, fertilization and seed development in this interesting group of plants. Various habitats and the economic importance of different species are also included. 29 minutes. CLEARVUE/eav, WW5VH 1614 Seedless Plants. The role that plants play as producers and how they contributed to the production of fossil fuel is explored, then how the seedless plants evolved for life on land is discussed. Structure and function of this group are also covered. 15 minutes. CLEARVUE/eav, WW5VH 1042 99