The first land plants: bryophytes ( non-vascular plants) 1. Traits shared by land plants, and lacking in the charophyceans
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1 Figure 29.1 Some highlights of plant evolution Figure 29.2x Chara Traits shared by charophyceans and land plants Very similar plastids (structurally similar, but especially similar chloroplast DNA) Very similar cellulose cell walls (cellulose is even produced by a similar rose-shaped structures) Anti-photorespiration enzymes packaged in peroxisomes Similar structure of flagellated sperm Similar structures during cell division (phragmoplasts) The first land plants: bryophytes ( non-vascular plants) 1. Traits shared by land plants, and lacking in the charophyceans 2. The earliest land plants: bryophytes (mosses, liverworts, hornworts) Figure 29.7 Land plant evolution. Figure 29.5 Land plant trait #1: Apical meristems, which are localized regions of cell division at the tips of shoots (left) and roots (right) 1
2 Figure 29.4 Land plant trait #2: Multicellular, dependent embryos, which develop from zygotes contained within tissues of the female parent. Parental tissues provide nutrients for the embryo. Land plants are also known as the embryophytes. Figure 29.6 Land plant trait #3: Alternation of generations life cycle, with two very different generations (one usually much larger & more prominent). Some algae also have alternation of generations, but not the charophyceans. Embryo Maternal tissue Figure 29.6 Land plant trait #4: Walled spores produced by a multicellular sporangium. A polymer called sporopollenin, the most durable organic material known, makes up the walls. Figure 29.5 Sporangium of a moss (a bryophyte) sporophyte Figure 29.5 Land plant trait #5: Gametangia, multicellular organs that produce gametes. Note: the most modern land plants, the flowering plants, do not have gametangia. Figure 29.9 Land plant trait #5: Gametangia, multicellular organs that produce gametes. Shown below are the Archegonium (egg-producing organ) of Marchantia (left), and the Antheridium (sperm-producing organ) of a hornwort (right). Note: the most modern land plants, the flowering plants, do not have gametangia. 2
3 Land plant trait #6: Specialized epidermis for water conservation, including a cuticle and stomata. Cuticle of a stem from Psilotum (a pteridophyte) is shown below. Figure Guard Cells regulate water loss through opening and closing of stomata. guard cell stomate Land plant trait #7: Adaptations for water transport, especially vascular tissue. Found in all land plants except for most bryophytes. The stem of Polypodium, a fern (a pteridophyte), is shown below. Note: true leaves, stems, and roots are defined by the presence of vascular tissues. Land plant trait #7: Adaptations for water transport, especially vascular tissue. Found in all land plants except for most bryophytes. The stem of Polypodium, a fern (a pteridophyte), is shown below. Note: true leaves, stems, and roots are defined by the presence of vascular tissues. true vascular tissues contain lignin Land plant trait #8: Secondary compounds: a diversity of chemicals with many functions related to living on land, including protection from UV radiation, signaling with symbiotic bacteria, deterring attack by pathogens or herbivores, and providing structural support. Quinine, a secondary compound produced by plants, is used by humans to help prevent malaria. Derived traits of land plants Apical meristems Plants move by growing Multicellular, dependent embryos thus Embryophytes Alternation of generations Heteromorphic 3
4 Derived traits of land plants Walled spores produced by multicellular sporangia Multicellular gametangia producing gametes Except in flowering plants (still have gametes) Spore vs. Gamete Spores grow directly into multicellular bodies Gametes need to undergo fertilization first Derived traits of land plants Epidermis for water conservation Cuticle, stomata Vascular tissues (except bryophytes) True vascular tissues have lignin True roots, stems, and leaves have vascular tissue Secondary compounds Protection, signaling, defense, support The first land plants: bryophytes ( non-vascular plants) Figure 29.1 Adaptation to living on land catalyzed a great deal of evolutionary diversification. Why? 1. Traits shared by land plants, and lacking in the charophyceans 2. The earliest land plants: bryophytes (mosses, liverworts, hornworts) Traits of bryophytes Figure 29.8 The life cycle of Polytrichum, a moss (Layer 3) Most lack true vascular tissue, which places limits on their thickness and height. In their alternation of generations life cycle, the gametophyte is the larger, conspicuous stage. The sporophyte is smaller, and when it grows, it is dependent on the gametophyte for nutrients. Liverworts have especially small sporophytes. Many mosses can live in extremely dry or cold habitats, because they can almost completely dry out without dying. 4
5 Figure 29.16x Stages of the moss life cycle. Key features of bryophyte life cycle Gametophyte is dominant generation Sporophyte is smaller, dependent on gametophyte (but still multicellular) Fertilization is dependent on free water carrying sperm from antheridia to archegonia Life Cycle of a bryophyte Life Cycle of a bryophyte Marchantia Polymorpha Marchantia Polymorpha Note the Male and female gametophytes Note the Male and female gametophytes Note the reduced sporophyte Note the reduced sporophyte Note antheridia and archegonia Note antheridia and archegonia Ecology of bryophytes Figure Sphagnum, or peat moss: Peat bog in Oneida County, Wisconsin (top), closeup of Sphagnum (bottom left), Sphagnum "leaf" (bottom right) All are photoautotrophs, primary producers Mostly live in moist habitats, although some mosses live in very cold and/or dry environments All need free water for fertilization, and this ultimately restricts their distribution 5
6 Figure 29.19x A peat moss bog in Norway. This landscape is probably similar to much of earth during the first 100 million years after land plants evolved: the only plants were short bryophytes. Why are bryophytes short? Peat (Sphagnum) bogs Cover nearly 1% of the Earth s land surface Dead peat resists decompostion, thus peat bogs store tremendous amounts of carbon (400 billion tons worldwide) Structure of peat moss is highly absorbent, making it useful to people Fuel Soil Conditioner Figure Sphagnum, or peat moss: Harvesting for human use Seedless Vascular Plants 1. Traits and adaptations of the first vascular plants: Pteridophytes 2. Two phyla of seedless vascular plants: Lycophyta (lycophytes) and Pterophyta (ferns & their relatives) 3. Overview of evolutionary transition to seed plants Figure 29.7 Land plant evolution. Figure 29.21x1 Lycophyte (club mosses) 6
7 Figure 29.21x2 Horsetail (Equisetum): Pterophyta Traits and adaptations of the first vascular plants Figure 29.23x1 A fern Figure Cooksonia, a vascular plant of the Silurian period. Note tall stature and large branched sporophyte with numerous sporangia. True lignified vascular tissue system (xylem and phloem), and thus true leaves, stems, and roots in most taxa Dominant sporophyte generation that is branched and becomes independent of the parental gametophyte No seeds Traits 1 & 2 are shared by the other two groups of vascular plants: gymnosperms and angiosperms Many adaptations to land inherited from bryophytelike ancestor (meristems, embryos, alt. of gen. life cycle, walled resistant spores, gametangia, specialized epidermis, secondary compounds) see previous lecture Figure Hypotheses for the evolution of leaves Microphylls, small (usually) leaves which have only a single strand of vascular tissue, are thought to have evolved from a small stem outgrowth or flap, into which a strand of vascular tissue grew. Megaphylls, large (usually) leaves with a branched vascular system, support more photosynthetic capacity than microphylls. They are thought to have evolved through the flattening and joining (via tissue webbing) of multiple branches. Rhizome (underground horizontal stem) Figure Hypotheses for the evolution of leaves Microphylls, small (usually) leaves which have only a single strand of vascular tissue, are thought to have evolved from a small stem outgrowth or flap, into which a strand of vascular tissue grew. Megaphylls, large (usually) leaves with a branched vascular system, support more photosynthetic capacity than microphylls. They are thought to have evolved through the flattening and joining (via tissue webbing) of multiple branches. 7
8 Note: some plants are homosporous and some are heterosporous. Sori Sporphylls - modified leaves that bear sporangia Strobili Homosporous Sporophyte Single type of spore Bisexual gametophyte (with both kinds of gametangia) Eggs & sperm Heterosporous Sporophyte Microspore Megaspore Male gametophyte Female gametophyte (inside the spore) Sperm Eggs The heterosporous life cycle was important in the evolution of the seed, which we ll discuss next week. Clusters of sporangia, usually on the underside of sporophylls Cones, usually on the tips of shoots/branches Seedless Vascular Plants 1. Traits and adaptations of the first vascular plants: Pteridophytes 2. Two phyla of seedless vascular plants: Lycophyta (lycophytes) and Pterophyta (ferns & their relatives) 3. Overview of evolutionary transition to seed plants Figure 29.7 Land plant evolution. Figure Pteridophytes (seedless vascular plants): Lycopodium (a club "moss, top left), Psilotum (a whisk fern, top right), Equisetum (a horsetail, bottom left), fern (bottom right). The latter three represent phylum Pterophyta, and Lycopodium represents phylum Lycophyta. Another genus in Lycophyta, which we will see in lab, is Selaginella. Figure Lycopodium (a club "moss ) in the phylum Lycophyta. Many grow on tropical trees as epiphytes Others grow on temperate forest floors Tiny gametophytes, sometimes underground Upright stems with many small leaves (microphylls) 8
9 Figure Selaginella (a spike "moss ) in the phylum Lycophyta. Figure Isoetes (a quillwort) in the phylum Lycophyta. Small, low to the ground Grow horizontally Represented today by a single genus Live in marshy areas Figure 29.7 Land plant evolution. Figure Equisetum (horsetail) in the phylum Pterophyta. Often have separate vegetative and fertile stems Jointed plants with rings of small leaves or branches at each joint Bulk of photosynthesis in stem Air canals carry oxygen to roots, which are often in waterlogged Soil Homosporous Modern Equisetum essentially identical to Equisetites from 300 mya. Equisetum may be the oldest surviving genus of plants on Earth. calamites were horsetails that grew into trees up to 18m tall and 45 cm thick during the Devonian (~300 mya) Figure Psilotum (whisk fern) in the phylum Pterophyta. Dichtomously branching stems, no roots (like first vascular plants) Stems have small scale-like outgrowths without vascular tissue - unclear if they predate leaves or are reduced leaves 9
10 Figure Polypodium (a fern) in the phylum Pterophyta. Figure The life cycle of a fern: note the dominant diploid (sporophyte) generation, and the reduced gametophyte FERNS ~12,000 species Most are in tropics Note megaphylls - fronds Most are homosporous Gametophytes shrivel and die Can produce LOTS of airborne spores Figure 29.23x2 Life cycle of a fern: sorus, a cluster of sporangia (~1 mm diam.) Figure 29.24c Closeup of a fern sorus. Note the sporangia. Figure 29.24b Underside of fern sporophyll, a leaf specialized for spore production. Note the numerous sori (clusters of sporangia) Figure 29.7 Land plant evolution. 10
11 Among the seedless plants: Figure Artist s conception of a Carboniferous forest, ~300 mya. Dead plants did not completely decompose, were later covered by seas and marine sediments, and then were transformed by heat and pressure into vast beds of coal. Mostly homosporous Spike mosses and quillworts (both Lycophytes) are heterosporous But, heterospory appears to have evolved independently in first seed plants (i.e., analogous), since seed plants appear more related to Pterophytes than Lycophytes Seedless Vascular Plants Figure 30.1 Three variations on gametophyte/sporophyte relationships. Note the progression in evolutionary time from left to right. 1. Traits and adaptations of the first vascular plants: Pteridophytes 2. Two phyla of seedless vascular plants: Lycophyta (lycophytes) and Pterophyta (ferns & their relatives) 3. Overview of evolutionary transition to seed plants Figure 30.3 From ovule to seed Advantages of the Seed Multicellular embryo gets a head start at the germination stage Stored food for embryo also allows a head start, and allows for extended dormancy Multicellular, larger and more complex than a spore = more resistant to harsh conditions Larger and more complex = increased capacity to develop dispersal adaptations 11
12 Disadvantage of the Seed Figure 30.2 From ovule to seed More energetically costly to produce To some extent, trade off quantity for quality Figure 30.3 Winged seed of a White Pine (Pinus strobus), a gymnosperm. Remember: Alternation of generations is thought to have evolved via delayed meiosis. What might be the advantage of this more complex life cycle, for living on land? Multicellular sporophyte = more than 4 spores produced by meiosis. Remember, algal sperm are flagellated, swim to eggs. This requires water. Harder to do on land. 12
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