Plant Diversity From Waterworld to Dry Land The Greening of Earth Looking at a lush landscape It is difficult to imagine the land without any plants or other organisms Figure 29.1 Plants Why Study Plants? Plants Provide Ecosystem Services Domestication and Selective Breeding Plant-Based Fuels and Fibers Bioprospecting How to Study Plants Analyzing Morphological Traits Using the Fossil Record Evaluating Molecular Phylogenies Themes in the Diversification of Plants The Transition to Land, I: How Did Plants Adapt to Dry Conditions? Preventing Water Loss: Cuticle and Stomata Transporting Water: Vascular Tissue and Upright Growth The Transition to Land, II: How Do Plants Reproduce in Dry Conditions? Retaining and Nourishing Offspring: Land Plants as Embryophytes The Evolution of Pollen The Evolution of the Seed The Evolution of the Flower The Angiosperm Radiation Why Study Plants? Among the most important endeavors supported by biological science Agriculture Forestry Horticulture Tens of thousands of biologists are employed in research designed to increase the productivity of plants and create new ways of using them that benefit people. Plants Provide Ecosystem Services Produce oxygen via oxygenic photosynthesis Build soil by providing food for decomposers Hold and prevent nutrients from being lost by erosion by wind and water Hold water Moderate the local climate. Plants are eaten by herbivores which are eaten by carnivores which are eaten by omnivores—organisms that eat both plants and animals. Omnivores feed at several different levels in the terrestrial food chain. Domestication and Selective Breeding Humans have actively selected seeds to plant the next generation of crops, a process called artificial selection Plant-Based Fuels and Fibers In addition to food, humans have depended on plants for cooking and heating fuels and as a source of fibers for clothing and other things Wood has been replaced by other fuels. Today, the primary interest in woody plants is for building materials and fibers used in papermaking. Bioprospecting The effort to find naturally occurring compounds Drugs Fragrances Insecticides Herbicides fungicides. Hydroponics The liquid culture of plants Can be used to harvest large quantities of plant chemicals. How Do Biologists Study Green Plants? To understand how green plants originated and diversified, biologists use three tools: 1. 2. 3. They compare the fundamental morphological features of various green algae and green plants; They analyze the fossil record of the lineage; and They assess similarities and differences in molecular traits such as the DNA sequences from selected genes Morphological Traits Important phyla of plants are grouped into three categories: 1. 2. 3. nonvascular plants seedless vascular plants seed plants Seed plants There are five major lineages in the group: 1. 2. 3. 4. 5. Cycads Ginkgoes Conifers Gnetophytes Collectively known as gymnosperms angiosperms Flowering plants Fossil Record The fossil record for land plants began 476 million years ago It is massive and is broken up into five segments, each of which encompasses a major event in the diversification of land plants Fossil Record Fossil Record Fossilized spores and tissues Have been extracted from 475-million-year-old rocks (a) Fossilized spores. Unlike the spores of most living plants, which are single grains, these spores found in Oman are in groups of four (left; one hidden) and two (right). (b) Fossilized Figure 29.6 a, b sporophyte tissue. The spores were embedded in tissue that appears to be from plants. Evaluating Molecular Phylogenies Points From Phylogenetic Tree Land plants probably evolved from green algae. The green algal group called Charales is the sister group to land 1. 2. • 3. Charales are their closest living relative. The green algae group is paraphyletic. Points From Phylogenetic Tree 4. 5. 6. 7. The land plants are monophyletic. The nonvascular plants are the most basal groups among land plants. Morphological simplicity of the whisk ferns is probably a derived trait. Seeds and flowers evolved only once. New Territory For more than the first 3 billion years of Earth’s history The terrestrial surface was lifeless Since colonizing land Plants have diversified into roughly 290,000 living species Plants Evolved from Green Algae Green algae have traditionally been considered protists, but we study them along with land plants for two reasons they are the closest living relatives to land plants the transition from aquatic to terrestrial life occurred when land plants evolved from green algae. charophyceans closest relatives Green Algae The green algae are a paraphyletic group that totals about 7000 species. They have a double membrane and chlorophylls a and b, but relatively few accessory pigments. Adaptations Enabling the Move to Land In charophyceans A layer of a durable polymer called sporopollenin prevents exposed zygotes from drying out The accumulation of traits that facilitated survival on land May have opened the way to its colonization by plants Derived Traits of Plants Five key traits appear in nearly all land plants but are absent in the charophyceans Apical meristems Alternation of generations Walled spores produced in sporangia Multicellular gametangia Multicellular dependent embryos Key Lineages of Green Plants Green Algae Ulvobionta Coleochaetales Charales (Stoneworts) Nonvascular Plants (“Bryophytes”) Hepaticophyta (Liverworts) Anthocerophyta (Hornworts) Bryophyta (Mosses) Seedless Vascular Plants Lycophyta (Lycophytes, or Club Mosses) Psilotophyta (Whisk Ferns) Sphenophyta (or Equisetophyta) (Horsetails) Pteridophyta (Ferns) Seed Plants Gnetophyta (Gnetophytes) Cycadophyta (Cycads) Ginkgophyta (Ginkgoes) Coniferophyta (Conifers) Anthophyta (Angiosperms) Nonvascular Plants, Bryophytes The most basal lineages of land plants Three lineages with living representatives Liverworts, phylum Hepatophyta Hornworts, phylum Anthocerophyta Mosses, phylum Bryophyta Not monophyletic Represent an evolutionary grade. Mosses are most closely related to vascular plants Bryophyte diversity Gametophore of female gametophyte LIVERWORTS (PHYLUM HEPATOPHYTA) Foot Plagiochila deltoidea, a “leafy” liverwort Seta Sporangium Marchantia polymorpha, a “thalloid” liverwort 500 µm Marchantia sporophyte (LM) HORNWORTS (PHYLUM ANTHOCEROPHYTA) An Anthoceros hornwort species Saprophyte MOSSES (PHYLUM BRYOPHYTA) Polytrichum commune, hairy-cap moss Sporophyte Gametophyte Figure 29.9 Gametophyte The life cycles of Bryophytes Gametophytes are larger and longer-living than sporophytes Bryophyte gametophytes Some mosses Have conducting tissues in the center of their “stems” and may grow vertically Bryophyte sporophytes Produce flagellated sperm in antheridia Produce ova in archegonia Generally form ground-hugging carpets and are at most only a few cells thick Grow out of archegonia Are the smallest and simplest of all extant plant groups Consist of a foot, a seta, and a sporangium Hornwort and moss sporophytes Have stomata Hepaticophyta, Liverworts Liver-shaped leaves can grow on bare rock or tree bark, which helps in soil formation Anthocerophyta, Hornworts The sporophytes look like horns and have stomata Bryophyta, Mosses May be abundant in extreme environments Can become dormant Sphagnum species are among the most profuse Seedless Vascular Plants Paraphyletic group Forms a grade between the nonvascular plants and the seed plants Have conducting tissues with cells that are reinforced with lignin, forming vascular tissue. The general groups of seedless vascular plants LYCOPHYTES (PHYLUM LYCOPHYTA) Strobili (clusters of sporophylls) Isoetes gunnii, a quillwort Selaginella apoda, a spike moss Diphasiastrum tristachyum, a club moss PTEROPHYTES (PHYLUM PTEROPHYTA) Psilotum nudum, a whisk fern Equisetum arvense, field horsetail Athyrium filix-femina, lady fern Vegetative stem Strobilus on fertile stem Figure 29.14 WHISK FERNS AND RELATIVES HORSETAILS FERNS Transport in Xylem and Phloem Xylem Conducts most of the water and minerals Includes dead cells called tracheids Phloem Distributes sugars, amino acids, and other organic products Consists of living cells Evolution of Roots Roots Are organs that anchor vascular plants Enable vascular plants to absorb water and nutrients from the soil May have evolved from subterranean stems Evolution of Leaves Leaves Are organs that increase the surface area of vascular plants, thereby capturing more solar energy for photosynthesis Leaves are categorized by two types Microphylls, leaves with a single vein Megaphylls, leaves with a highly branched vascular system One Model of Evolution Microphylls evolved first, as outgrowths of stems Vascular tissue Figure 29.13a, b (a) Microphylls, such as those of lycophytes, may have originated as small stem outgrowths supported by single, unbranched strands of vascular tissue. (b) Megaphylls, which have branched vascular systems, may have evolved by the fusion of branched stems. Phylum Lycophyta: Club Mosses, Spike Mosses, and Quillworts Modern species of lycophytes Are relics from a far more eminent past Are small herbaceous plants Lycophyta, Club Mosses Most ancient plant lineage with roots Tree-sized dominated the coal-forming forests of the Carboniferous period. Psilotophyta, Whisk Ferns Restricted to tropical regions Have no fossil record Sphenophyta (or Equisetophyta), Horsetails Flourish in waterlogged soils by allowing oxygen to diffuse down their hollow stems Pteridophyta, Ferns only seedless vascular plants to have large, well-developed leaves