Tuesday, November 20, Plants Invade the Land

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1 Plants Invade the Land

2 ?

3 What is a plant?

4 What is a plant? A nonmotile, desiccation-resistant, structurally reinforced, photosynthetic organism with cellulosic walls.

5 What is a plant? A nonmotile, desiccation-resistant, structurally reinforced, photosynthetic organism with cellulosic walls. Plant cell Animal cell

6 What is a plant? A nonmotile, desiccation-resistant, structurally reinforced, photosynthetic organism with cellulosic walls.

7 What is a plant? Cannabis A nonmotile, desiccation-resistant, structurally reinforced, photosynthetic organism with cellulosic walls. Opium poppy Coffee Nicotine Peyote Coca Datura (for making zombies) Theobroma (chocolate)

8 What is a plant? A nonmotile, desiccation-resistant, structurally reinforced, photosynthetic organism with cellulosic walls. Perforation plate (compound) Substomatal cavity Palisade parenchyma Xylem Perforation plate (simple) Cuticle Upper epidermis Air boundary layer Mesophyll cells High water vapor content Low CO 2 Lower epidermis Cuticle Boundary layer resistance (r b ) Water vapor Low water vapor content Leaf stomatal resistance (r s ) CO 2 High CO 2 Guard cell Stomatal pore Air boundary layer Pits Vessel elements FIGURE 4.10 Water pathway through the leaf. Water is pulled from the xylem into the cell walls of the mesophyll, where it evaporates into the air spaces within the

9 What is a plant? A nonmotile, desiccation-resistant, structurally reinforced, photosynthetic organism with cellulosic walls. End wall of vessel element with bordered pits Gas-filled cavitated vessel Scalariform perforation plate Pit Liquid water Water vapor bubble Gas-filled cavitated tracheid P-protein Sieve tube element Modified plastid Sieve tube element Smooth endoplasmic reticulum Cytoplasm Plasma membrane Thickened primary wall elements (sieve tube elem showing sieve plates and Longitudinal section, show ments joined together to f pores in the sieve plates b ments are open channels f sieve tube. The plasma me element is continuous wit sieve tube element. Each s ciated with one or more c take over some of the esse that are reduced or lost du sieve tube elements. Note has many cytoplasmic org tube element has relativel nary companion cell is de Companion cell Branched plasmodesmata Vacuole Chloroplast Nucleus Secondary phloem Vascular cambium Pith Secondary xylem Sieve plate pore Mitochondrion Sieve plate

10 What is a plant? A nonmotile, desiccation-resistant, structurally reinforced, photosynthetic organism with cellulosic walls. Lignin Wood

11 What is a plant? A nonmotile, desiccation-resistant, structurally reinforced, photosynthetic organism with cellulosic walls. Total solar energy (100%) H 2 O ADP + P i NADP + (CH 2 O) n Light Chlorophyll Triose phosphates Nonabsorbed wavelengths (60% loss) O 2 ATP + NADPH CO 2 + H 2 O 40% Light reactions Carbon reactions 32% Reflection and transmission (8% loss) Heat dissipation (8% loss) 24% Metabolism (19% loss) 5% Carbohydrate

12 What is a plant?

13 What is a plant? fossil

14 Stems What is a plant? fossil Spores and cones Rhynia Medullosa Horneophyton spores Leaves and branches Conifer cone Alethopteris Conifer cone Sawdonia Neuropteris

15 Photosynthesis on Land

16 Photosynthesis on Land

17 Photosynthesis on Land Photosynthesis is fundamentally aquatic: 1 cm Acetabularia Halimeda The atmosphere is really dry! Dangers of living on land: No more currents to aid dispersal! Can t get CO 2 without losing water! No more buoyancy to help stay up! How do land plants cope?

18 Photosynthesis on Land The atmosphere is really dry! Can t get CO 2 without losing water! Protect spores with sporopollenin te Cuticle keeps leaves and stems from drying out; Lignin allows plants to grow tall; Stomata are pores that regulate CO 2 and H 2 O exchange: (singular: stomate not stoma )

19 Photosynthesis on Land No more currents to aid dispersal! No more buoyancy to help stay up!

20 Photosynthesis on Land Grow tall! No more currents to aid dispersal! No more buoyancy to help stay up!

21 Photosynthesis on Land Grow tall! No more currents to aid dispersal! No more buoyancy to help stay up! Archegonia of Marchantia, a liverwort

22

23 ANGIOSPERMS >250,000 species GYMNOSPERMS 854 species FERNS ~10,000 species LYCOPHYTES ~1,250 species BRYOPHYTES ~25,000 species

24 Laminar leaves Wood Seeds Flowers Vascular tissue Marine Terrestrial

25 Lycopod Trees Seeds Wood Seeds Wood Medullosa?? Rhyniophytes Cladoxylopsids Flowers Laminar leaves Laminar leaves Wood Seeds Vascular tissue Marine Terrestrial

26 Plants Invade the Land First unambiguous spores: 475Ma, Libya First unambiguous sporophytes: ~430Ma, Cooksonia First seeds: ~390Ma, West Virginia, Elkinsia 20 microns 20 microns From Wellman et al., Nature 2003: Fragments of the Earliest Land Plants 1 cm 1 cm 35 m First trees: ~380Ma, Archaeopteris Cooksonia Elkinsia Archaeopteris

27 Terrestrial Pioneers Typical Ordovician and Silurian spores Distribution of Ordovician terrestrial spores (Gensel et al., Plants Invade the Land)

28 Trees with true roots Seeds Trees Plants taller than 2-3cm Terrestrial plant spores From Kenrick and Crane, Nature 1997: Origin and Evolution of Plants on Land

29 The Rhynie Chert William Mackie Discovered in 1912, described in by Kidston and Lang Age: 404 ± 2 Ma Volcanic sinter/geyser complex (like Yellowstone)

30 The Rhynie Chert Unpolished hand sample Nonvascular plants and their spores Vascular plants, preserved in place, with small leaves!

31 Rhynie Chert Flora

32 The Rhynie Chert By the time of the Rhynie Chert, we already see representatives from these two groups and some fungi and spiders!

33 The Rhynie Chert Symbiotic fungi (mycorrhizae) Webless arachnid inside a plant stem

34 Middle Devonian Flora: Vascular plants Ancestors of seed plants and ferns Lycophytes Nonvascular plants 10 cm Middle Devonian, ~400Ma: tiny plants

35 M-L Devonian: More Diversity Increase in ecosystem complexity and rooting depth through the Devonian. Effects on biogeochemical cycles? Some odd-looking Devonian trees (the cladoxylopsids) Archaeopteris, the first tree

36 M-L Devonian: More Diversity Increase in ecosystem complexity and rooting depth through the Devonian. Effects on biogeochemical cycles? Some odd-looking Devonian trees (the cladoxylopsids) Archaeopteris, the first tree

37 The Age of Coal

38 Arborescent Lycopods 30m-tall trees with very little wood and branching! Tree-sized herbs : grew fast, didn t live long. Dominated swampy, black-water environments.

39 Coal Swamp Ecology Coal lycopod trees Shale seed plants and ferns Paleo-river channel preserved in Illinois Seed plants on river banks lycopods in swamps!

40 The Seed Plant Revolution Seed plants (Some) Cladoxylopsids Ferns Calamopityales Medullosans Ginkgo Cordaites Conifers Glossopteris Angiosperms 240 Ma 318 Ma 310 Ma 125 Ma Laminar Leaves Laminar Leaves 320 Ma? 300 Ma? 385 Ma 359 Ma 350 Ma?Laminar leaves 390 Ma Extinct taxa Extant taxa Poly/paraphyletic taxa Seed plants originate on river banks and in the forest understory, but don t run the swamps until the Mesozoic!

41 In 50 million years, from this: Vascular plants Ancestors of seed plants and ferns Lycophytes Nonvascular plants 10 cm Middle Devonian, ~400Ma: tiny plants

42 to this: Giant dragonfly Seed plants Lycophytes Ferns 1 m Carboniferous swamp forests, ~350Ma

43 Late Paleozoic Biogeography The dominant plants of the Mesozoic first appear in the Paleozoic tropics

44 Mesozoic Plants Ginkgo biloba Tree Ferns Cycads

45 Mesozoic Plants Ginkgo biloba Tree Ferns David Bowie: famed cycad collector, also a musician Cycads

46 Mesozoic Plant Diversity Triassic Conifers First collected by W. T. Sherman, 1879

47 Roots of Modern Flora K/T impact

48 Flowering Plant Story: Fossils ~80 million-year-old mummified flower, Georgia (USA) (Crane et al. 2004) Cretaceous angiosperm leaves ~90 million-year-old mummified flowers, Maryland (USA) Mesofossils! Archaefructus, a Cretaceous angiosperm

49 K T Angiosperm diversity is rooted in the Cretaceous...

50 80 Angiosperms also displace seed plants from many environments. but where did they come from? Mean number of species per flora Other seed plants Cordaites Caytonioids Cycadeoids Angiosperms 20 Sphenopsids Conifers 10 Trimerophytes Lycophytes Ferns Pteridosperms Cycads P Progym Z R! Ginkgos S Devonian Miss. Penn. Perm. Triassic Jurassic Cretaceous Tertiary Age (Ma) Redrawn from Knoll (1986) (using original age and taxonomic assignments)

51 Welwitschia, only found in Southern Africa Ephedra, found in Asia, N. & S. America, diet stores Gnetum, found in SE Asia (tropical forests)

52 First cycads are nearly 300 million years old......where are the Paleozoic flowers?

53 Candidates? Clockwise from above: Cycadeoidea, a Mesozoic cycadlike plant; Williamsonia, a quasibisporangiate relative of Cycadeoidea; extant Welwitschia; Glossopteris, Paleozoic seed plant; Caytonia, another Mesozoic seed plant.

54 Conclusions Plants required a complex suite of biophysical adaptations to live on land. Virtually all of these adaptations appeared within 50 million years of the first terrestrial plants. Paleozoic forests had a remarkably different taxonomic composition from modern forests. Everything you eat, wear, and use is the product of the Cenozoic radiation of flowering plants.

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