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Vascular Plants. Generalized life cycle. Moss life cycle. Spore -> Gametophyte. Gametophyte -> Zygote. Zygote -> mature sporophyte. Sporophyte -> spores. Vascular Plants. Vascular Plants. Branching sporophyte in Silurian First vascular plants in Devonian. First flowering plants. - PowerPoint PPT Presentation
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Vascular Plants
Generalized life cycle
Moss life cycle
Spore -> Gametophyte
Gametophyte -> Zygote
Zygote -> mature sporophyte
Sporophyte -> spores
Vascular Plants
Vascular Plants
• Branching sporophyte in
Silurian• First vascular plants in Devonian
Liverwort spore tetrads - end of Ordovician First fossil of non-algal land plant in Silurian - 430 mya
First flowering plants
Age of Ferns
Fossils• Probable embryophyte spores at 450 Ma
• Cooksonia fossils in Silurian (ca. 430 Ma)
Plants like Cooksonia lacked a vascular system
• Dichotomously-branching axes
• Terminal sporangia• No roots or leaves
Over-time became larger, more complex, and acquired
a vascular system
Time
Living vascular plants
(Tracheophytes)• Stems and roots (often leaves)
Living vascular plants (Tracheophytes)
• Stems and roots • Sporophytes dominate the life-cycle
Tree fernTree fern
Sporophyte Sporophyte dominancedominance
Gametphyte
Why sporophyte dominance?
• Spore dispersal by wind: aided by height
• Competition for light (gametophyte constrained by the need for water)
Living vascular plants (Tracheophytes)
• Stems and roots • Sporophytes dominate the life-cycle
• A vascular (transport) system
Vascular system• Xylem (water transport) and phloem (metabolite transport)
Xylem Phloem QuickTime™ and aTIFF (Uncompressed) decompressor
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Stem
Root
Vascular Bundle
Xylem
• Composed primarily of Tracheids
• Elongated, dead, cells
• Cell wall impregnated with lignin
• Transport of water from soil to leaves
Phloem• Transport via Transport via sieve elementssieve elements
• Elongated, Elongated, living cellsliving cells
• Transport of Transport of sugars, sugars, hormones, etc.hormones, etc.
3 Major groups of Vascular plants
• Seed plants (gymnosperms
and angiosperms)
• Lycophytes (club mosses
and their relatives)
• Moniliforms (ferns and
fern allies)
Major tracheophyte taxa
• Seed Plants (ca. 290,000 species)
• Lycophytes (ca. 1,100 species)
• Ferns and allies (ca. 11,000 species)Tracheophytes that are not Tracheophytes that are not seed plants are sometimes seed plants are sometimes called “pteridophytes”called “pteridophytes”
Vascular plant phylogenybryophytes
Cooksonia
lycophytes Ferns + Seed plants
Branched sporophyte
Vascular tissue,Roots
Megaphylls
Microphylls
Organ systems origins
• Stems - dichotomous branching
Organ systems origins
• Stems• Leaves
– Microphylls
Organ systems origins
• Stems• Leaves
– Microphylls– Megaphylls
Organ systems origins
• Stems• Leaves
– Microphylls– Megaphylls
• Roots
Homospory versus Heterospory
An important variation: Heterospory
Selaginella (heterosporous)
MegasporangiumMegasporangium
MicrosporangiumMicrosporangium
Lycopodium(homosporous)
Heterospory• Microspores and megaspores produced in different sporangia on different leaves (microsporophylls; megasporophylls)
• Microspores grow into male gametophytes
• Megaspores grow into female gametophytes - remains within spore wall
Haploid
Diploid
Homospory
gametophyte
sperm
eggzygote
sporophyte
spore
HaploidDiploid
Heterospory
femalegametophyte
sperm
eggzygote
sporophyte
megasporemicrospore
malegametophyte
Heterospory evolved many times. Why?
• Increases potential for outcrossing
• Specialization of function between micro- and megagametophyte permits greater efficiency (less cost)
3 Major groups of Vascular plants
• Seed plants (gymnosperms
and angiosperms)
• Lycophytes (club mosses
and their relatives)
• Moniliforms (ferns and
fern allies)
Moniliforms
Spermatophyta
Lycophyta
Pryer et al. 2001
Lycophytes
• 380 Ma old• 1100 spp.• Microphylls only• Sister group to the other living vascular plants
Lycophytes
• 380 Ma old• 1100 spp.• Microphylls only
lycophytes Ferns + Seed plants
Megaphylls
Microphylls
Lycophytes
LycopodiumLycopodium
SelaginellaSelaginella Isoetes
Selaginella
Clubmoss (Lycopodium)
LepidodendronLepidodendronSigillariaSigillaria
Carboniferous lycopodsup to 40 m
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Lycopodium life cycle I
Sporophyte makes sporangia often in a “strobilus” Sporangium
Lycopodium life cycle I
Sporophyte makes sporangia often in a “strobilus”Sporophyll
Sporangium
Lycopodium life cycle II
Spores dispersed by wind: germinate into a minute gametophyte
Spore Thallus
Rhizoids
Spore
Lycopodium life cycle III
• Gametophyte produces archegonia and antheridia (bisexual)
• Biflagellate sperm fertilize egg cells
• New sporophyte grows
Archegonial neck
Lycophyte diversity
• 3 Major groups– Lycopodiaceae (club mosses)
– Selaginella– Isoetes
Lycopiaceae• Approximately 400 species
• Dominated Carboniferous, up to 40 m tall– form much of modern coal
• Homosporous• Archegonia and Antheridia can take 6-15 years to mature
Selaginella• Approximately 700 extant species
• Heterosporous• Moist habitats or "resurrect"
Selaginella umbrosa
Isoetes• Approximately 200 species
• Grow in water or dried pools
Moniliforms
Spermatophyta
Lycophyta
Ferns and fern allies (moniliforms)
• Includes ferns (Pterophyta) and two small groups (Psilophyta and Sphenophyta)
• ca. 12,000 spp.• Homosporous or heterosporous• Megaphylls (lost in Psilophyta and Sphenophyta)
• Ecologically important especially as tropical epiphytes
Moniliforms
Ferns and Fern Allies
Ferns and Fern Allies
Psilotum
Equisetum
Ferns and Fern Allies
eusporangiate ferns
Sporangium wall has 2 or more cell layers
Ferns and Fern Allies
leptosporangiate ferns
Sporangium wall has 1 cell layer
Ferns and Fern Allies
Psilotum and Ophioglossum
Psilotaceae/Psilophyta
• Psilotum (2 spp.) and Tmesipteris (15 spp.)
• No roots and reduced or absent leaves, photosynthetic stems
• Sporangia on lateral branches
• Homosporous
• Thought to be "primitive vascular plants", but more likely simplified due to association with fungi.
Psilotaceae
Sporangium
Reduced forked leaves
Tmesipteris
Psilotum
Psilotum
Psilotum
Dichotomizing stem, no roots Long-lived gametophytes
OphioglossaceaeAdder's tongue, eusporangiate ferns
• Homosporous• Worldwide, common in disturbed areas
• Botrychium (~60 spp.) and Ophioglossum (30 spp.)
• Ophioglossum can have upwards of 1400 chromosomes - perhaps more then any other organism
Ophioglossaceae
Sterile blade
spore bearing sporophore
OphioglossaceaeBotrychium virginianum
Ferns and Fern Allies
Equisetum
EquisetaceaeHorse tails
• Equisetum (15 spp.)• Homosporous• Dates back to Devonian, with 20 m high stems - lots of diversity in Carboniferous forests
• Extant species "living fossils"• Leaves whorled, fused into sheaths at base, only microphylls
Equisetum
Calamites (Carboniferous)
Extinct trees
Calamites
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Equisetum gametophyte (hermaphroditic)
leptosporangiate ferns
Osmunda
Most ferns species are Leptosporangiate Ferns
Large megaphylls (fronds) unfold lengthwise from a "fiddlehead"
Section through sorus
Sporangium
Sporangia are arranged in sori
SporangiaIndusium
Sori
Fern gametophyte (prothallus)
Ferns Walking fern
Epiphytic fern (Platycerium)
Maidenhair fern Tree ferns (Cyatheaceae)
Leptosporangiate ferns
• App. 11,000 species in 25-35 recognized groups
(most of fern diversity)• Cover 4 of the major clades
- Marsileaceae- Osmundaceae- Cyatheaceae- Polypodiaceae
Marsileaceae
SalviniaSalvinia MarsileaMarsilea
• Mostly aquatic• Leave blade divided into 2-4 leaflets (clover-like)• Heterosporous
- megagametophytes with only one archegonium
Symbiotic with cyanobacteria, fertilized rice fields
Spores remain viable for a century
Osmundaceae
Sporangia loose, not in soriHomosporous
Osmunda
Leptopteris
Todea
Cyatheaceae
Tree fern growthSporangia in sori on bottom of leafStem usually single and erect
Alsophila
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Cyathea
PolypodiaceaeAsplenium
Adiantum
Pteris
Polypodium
Tectaria
Nephrolepis Elaphoglossum
Main points
• Features of vascular plants• Homospory versus heterospory• Megaphylls vs. microphylls • Life cycle of the fern• Fern allies: Psilotum, Equisetum
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