EVOLUTION and Diversity of GREEN and LAND PLANTS UNIT II EVOLUTION and DIVERSITY of PLANTS 59

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EVOLUTION and Diversity of GREEN and LAND PLANTS UNIT II EVOLUTION and DIVERSITY of PLANTS 59 3 EVOLUTION AND DIVERSITY OF GREEN AND LAND PLANTS THE GREEN PLANTS 55 Hornworts 69 70 EMBRYOPHYTA- LAND PLANTS 59 Polysporangiophytes/Pan-Tracheophyta DIVERSITY OF NONVASCULAR LAND PLANTS 62 REVIEW QjESTIONS 71 Liverworts 62 EXERCISES 72 Mosses 65 REFERENCES FOR FURTHER STUDY 72 THE GREEN PLANTS the cells, acting as a sort of cellular exoskeleton. The evolu tion of a cellulosic cell wall was a preamble to the further The green plants, formally called the Viridiplantae or evolution of more complex types of growth, particularly of Chiorobionta, are a monophyletic group of eukaryotic organ self-supporting shoot systems. It is not clear if a cellulosic isms that includes what have traditionally been called “green cell wall constitutes an apomorphy for the Viridiplantae algae” plus the land plants or embryophytes (Figure 3.1). alone, as it may have evolved much earlier, constituting an Like all eukaryotes, the Viridiplantae have cells with apomorphy for the Viridiplantae plus one or more other membrane-bound organelles, including a nucleus (containing groups; in any case, its adaptive significance seems clear. chromosomes composed of linear chains of DNA bound to Perhaps the primary apomorphy for the Viridiplantae is a proteins, that are sorted during cell division by mitosis), micro specialized type of chloroplast (Figure 3.2). As discussed in tubules, mitochondria, an endoplasmic reticulum, vesicles, Chapter 1, chloroplasts are one of the major defining charac and golgi bodies. Although the interrelationships of the non— teristics of traditionally defined “plants”; their adaptive sig land plant Viridiplantae will not be covered in detail here, it nificance as organelles functioning in photosynthesis, the is important to realize that some of the evolutionary innova conversion - of light energy to chemical energy, is unques tions, or apomorphies, that we normally associate with land tioned. Chloroplasts in the Viridiplantae, the green plants, differ plants actually arose before plants colonized the land. from those of most other organisms, such as the red and brown Several apomorphies unite the Viridiplantae (Figure 3.1). “algae,” in (1) containing chlorophyll b in addition to chloro One possible novelty for this group is a cellulosic cell wall phyll a, the former of which acts as an accessory pigment in (Figure 3.2A). Cellulose, like starch, is a polysaccharide, light capture; (2) having thylakoids, the chlorophyll-containing but one in which the glucose sugar units are bonded in the membranes, that are stacked into grana, which are pancake- beta-l,4 position (=3-1,4-g1ucopyranoside). This slight change like aggregations (see Figure 3.2B,C); and (3) manufacturing as in chemical bond position results in a very different molecule. a storage product true starch, a polymer of glucose sugar Cellulose is secreted outside the plasma membrane as micro units (= polysaccharide) in which the glucose molecules are scopic fiber-like units called microfibrils that are further chemically bonded in the alpha-1,4 position (ci-l,4- intertwined into larger fibril units, forming a supportive glucopyranoside). Thus, all green plants, from filamentous meshwork. The function of cellulose is to impart rigidity to green “algae” in a pond or tide pooi to giant sequoia or 55 02010 Elsevier Inc. All rights reserved. doi: 10. 1016/B978-0- 12-374380-0.00003-9 r 56 CHAPTER 3 EVOLUTION AND DIVERSITY OF GREEN AND LAND PLANTS UNIT II EVOLUTION AND DIVERSITY OF PLANTS 57 Viridiplantae [Chiorobionta] — Green Plants Chiorophytes 1i Streptophytes “Green Algae” (a paraphyletic group) Charophytes -IE Land Plants fi = Embryophytes FIGURE 3.3 Examples of non—land plant Viridiplantae. A. Chlansydomonas reinhardtii, a unicellular form. (Photo courtesy of archegonium Rick Bizzoco.) B. Ulva, a thalloid form. C. Volvox, a colonial form. D. Spirogyra, a filamentous form. Above: vegetative form, with large, chloroplasts. antheridium spiral Below: reproductive conjugation stage, showing + and — mating strains and nonmotile zygotes. parenchyma cuticle Eucalyptus trees have this same type of chioroplast. Recent cells in at least one phase of their life history. “Green data imply that chioroplasts found in the green plants today algae” inhabit fresh and marine waters sporophyte/embryo (alternation of generations) and some live in or on were modified from those that evolved via endosymbiosis, soil (or even on snow!) or in other terrestrial but moist the intracellular cohabitation of an independently living, uni habitats. cellular prokaryote inside a eukaryotic cell (see Chapter 1). The primitive type of green plant sexual reproduction The Viridiplantae as a whole are classified as two sister seems to have been the production of flagellate, haploid (n) groups: chiorophytes, or Chlorophyceae, and streptophytes, gametes that are “isomorphic,” that is, that look identical. or Streptophyceae true starch storage compound (Figure 3.1). The traditional “green algae” Fertilization occurs by union of two of these gametes, result are a paraphyletic group (which is why the name is placed in thylakoids stacked in grana Unique green plant chloroplast ing in a diploid (2n) zygote (Figure 3.4A). The zygote, which features quotation chlorophyll b (chlorophyll a is ancestral) marks) and are defined as the primarily aquatic is free-living, then divides by meiosis to form four haploid Viridiplantae, cellulose in cell wall (may have evolved consisting of all chiorophytes and the non—land spores, each of which may germinate and develop into a new earlier & thus not a synapomorphy for Chlorobionta alone) plant streptophytes. “Green algae” occur in a tremendous haploid individual, which produces more gametes, complet FIGURE variety of morphological 3.1 Cladogram of the green plants (Viridiplantae or Chlorobionta), forms. These include single cells ing what is termed a haplontic (or “haplobiontic”) life cycle modified from Bremer (1985), Mishler and Churchill (1985), and Mishler et al. (1994). Important apomorphies discussed (Figure 3.3A) with or without flagella, thalloid forms (Figure (Figure 3.4A). in the text are listed beside thick hash marks. 3.3B), motile and nonmotile colonies (Figure 3.3C), and non- Within the streptophyte lineage that gave rise to the land motile filaments (Figure 3.3D). Many have flagellated motile plants, a few innovations evolved that may have been HAPLOID (n) HAPLOJD (n) //Z Multicelled Stage Multicelled Stage granum — ..- - .thylakoid -7:-’ : mitosis I / HAPLONTIC •; •; C HAPLONTIC . / Isogamy Oogamy Spores Gamete Gamete Spores i Egg Sperm (n) 1/(n) (n) (n) \ (n) (n) fertilization // -i’‘i•: JneiosisZ3ote nze,oszs fertilization FIGURE Zygote 3.2 A. Elodea, whole leaf in face view, A showing apomorphies of the Viridiplantae: a cellulosic chloroplasts. B. Diagram cell wall and green plant of chloroplast structure of green plants, showing (2n) (2n) thylakoids and grana. C. Electron micrograph of Chlamydomonas reinhardtii, a unicellular “green alga,” showing granum of chioroplast. (Photo courtesy of Rick Bizzoco.) FIGURE 3.4 Haplontic life cycles in some of the green plants. A. Jsogamy. B. Oogamy. ii 56 CHAPTER 3 EVOLUTION AND DIVERSITY OF GREEN AND LAND PLANTS UNIT II EVOLUTIONANDDIVERSITYOFPLANT5 57 Viridiplantae [Chiorobionta] — Green Plants Chiorophytes r —j Streptophytes “Green Algae” (a paraphyletic group) J:: Charophytes - - . Land Plants = Embryophytes I FIGURE 3.3 Examples of non—land plant Viridiplantae. A. Chiarnydornonas reinhardtii, a unicellular form. (Photo courtesy of archegonium Rick Bizzoco.) B. Ulva, a thalloid form. C. Volvox, a colonial form. D. Spirogyra, a filamentous form. Above: vegetative form, with large, antheridjum spiral chloroplasts. Below: reproductive conjugation stage, showing + and — mating strains and nonmotile zygotes. parenchyma cuticle Eucalyptus trees have this same type of chloroplast. Recent cells in at least one phase of their life history. “Green data imply that chioroplasts found in the green plants today algae” inhabit fresh and marine waters and sporophyte/embryo (alternation of generations) some live in or on were modified from those that evolved via endosymbiosis, soil (or even on snow!) or in other terrestrial but moist the intracellular cohabitation of an independently living, uni habitats. cellular prokaryote inside a eukaryotic cell (see Chapter 1). The primitive type of green plant sexual reproduction The Viridiplantae as a whole are classified as two sister seems to have been the production of flagellate, haploid (n) groups: chiorophytes, or Chlorophyceae, and streptophytes, gametes that are “isomorphic,” that is, that look identical. true starch storage compound or Streptophyceae (Figure 3.1). The traditional “green algae” Fertilization occurs by union of two of these gametes, result thylakoids stacked in grana are a paraphyletic group (which is why the name is placed in ing in a diploid (2n) zygote Unique green plant chioroplast features (Figure 3.4A). The zygote, which chlorophyll b (chlorophyll a is ancestral) quotation marks) and are defined as the primarily aquatic is free-living, then divides by meiosis to form four haploid cellulose in cell Viridiplantae, wall (may have evolved earlier & thus consisting of all chiorophytes and the non—land spores, each of which may germinate and develop into a new not a synapomorphy for Chiorobionta alone) plant streptophytes. “Green algae” occur in a tremendous haploid individual, which produces more gametes, complet FIGURE 3.1 Cladogram of the green variety of morphological
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